American Fern Journal A QUARTERLY DEVOTED TO FERNS Published by the AMERICAN FERN SOCIETY e Epirors DAVID B. LELLINGER C. V. MORTON ROLLA M. TRYON IRA L. WIGGINS MONUMENTAL PRINTING COMPANY, BALTIMORE, MARYLAND Missourt BOTANICAL GARDEN LIBRARY Contents VotuME 60, NuMBER 1, Paczs 1-48, IssuED Aprix 21, 1970 Aberrant Leaves on Angle-Shoots of Selaginella martensii Spring Terry R. Webster A Further Note on the Type of Platycerium alcicorne..... C. V. Morton Gametophyte Ontogeny and Sex Expression in Dryopteris ludoviciana tchael I. Cousens and Harry T. Horner Shorter Notes: A New Form of Microlepia speluncae; Two New State Records for Isoétes in Kentucky; Psilotum nudum New to Arizona... PMNS AL OC a ee Re aay American Fern Society: Report of the President; Report of the Secretary; Report of the Treasurer; Report of the Auditing Committee; Report of the Judge of Elections; Report of the Fern Spore Exchange VoLuME 60, NUMBER 2, Paces 49-88, IssuED Juy 3, 1970 The Fern Collections in Some European Herbaria, VI... . . C. V. Morton Fertile Sporophytes of Botrychium virginianum Attached to Gameto- phytes Pepe Fane Se Sigs te ie ck Coe OS Bruce W. McAlpin A Peculiar Species of Grammitis:..... 2.2... 0. C. V. Morton Vascular Organization of the Rhizome of Cibotium barometz Subhash Chandra A Survey of Some Morphological Features of the Genus Elaphoglossum ee a ee Robert M. Lloyd Notes and News Sa Se ra phyt Cystopteris tenuisecta........ S. S. Bir i . ici vestigation of the Mexpholeny. of aoe eben! crenatum se saad and Nisha Bajpat A Major North American Range Extension for the Forked Spleenwort, Aspleni septentrionale David L. Emory Hard Water as a Limiting Factor i in the Distribution of Isoétes echino- Le e and Frances Belknap t Notes: Stem | Cross-section Prints Aid in eh eae Horsetails; dum - Northward; Terminology of the Spor- 8 -angial Structures of Equisetum; Kinds of Cystopteris in bate Vou. 60 JANUARY—Marcu, 1970 No. 1 American Fern Journal A QUARTERLY DEVOTED TO FERNS Published by the AMERICAN FERN SOCIETY ts DAVID B, LELLINGER Cc. V. MORTON ROLLA M. TRYON IRA L. WIGGINS Misaoue: Bora Nicac CONTENTS GanDed Lorany, etenenetens Te — es Aberrant Leaves on Angle-Shoots of Selaginella martensii Spring Terry R. Wesstsr 1 A Further Note on the —— of Platycerium alcicorne.....C. V. Morton 7° r' pt y A Ontogen Tt. %, r | MicHarEL tt CovusENS AND Harry T. Horner 13 Shorter Notes: A New Form of Microlepia speluncae; Two New State Records for Isoétes in Kentucky; Psilotum nudum New 28 Notes and News............. ae 32 Detedt Varn Liieenbne: oo re Ci a a eee American TT a La y Tw p -« 2 ait. kT. a wp p £ at a . vs R rt of th ss Rep + of th A | gC itt: ; Report of a. v. 1 40 = th ae re D1 a: D ‘er ye ee, e Judg ; The American Fern Hocietp Council for 1970 Warren H. ——— JR., —— of Botany, University of Michigan, 481 Joun T. Theis New York Botanical Garden, Bronx Park, Bronx, New 0458. ice-President Murray Evans, Department of Botany, University of ‘Toman, Knox- ville, Tenn. 37916. cretary LeRoy K. Henry, Division of Plants, Carnegie Museum, Pittsburgh, Penn- atvenie 15213. easurer Davip B. Letuincer, Smithsonian Institution, Washington, D. C. 20560. Editor-in-Chief Fiational Societp Representatives Warren H. Wacner, JR., oe of — A.A.A.S. Representative Rouiia M. Tryon, Jr., Harvard Uni A.1.B.S. Representative American = Journal EDITORS Davi B. LELLINGER..... Smithsonian Institution, Washington, D. C. 20560. ee VW, MORTON, ceca cess Smithsonian Institution, Washington, D. c 20560. LLA Pattie, Harvard University, Cambridge, Mass. 02138 Tra L. Wiaors.. . Dudley Herbarium, Stanford University, Stanford, Calif. An illustrated quarterly devoted to the general study of “la owned by Fern Society, and pid at Be at 3110 Ave., Baltimore those interested in obta’ ci Matter = uliation sheekd bs be addressed to the Editor-in-Chi dress, = cations for membership, “subscriptions, orders uld be addressed to the Subscriptions $5.50 gross, $5.00 net (agency fee $0. 50); sent ~ to mem- American Fern Rates Ager al dues, $5.00; sustaining membership, i ). Extracted re reprints, if dent ji . advance, at cost, plus postage. a eng pace Foo peg single back numbers of 64 Gummuative less, $1 pages, $2.00 each; sorte SD peep, $2.50 each; tive Tndex to Volumes 225, 1-25, 50 cents. Ten Librarp and Berbarium Dr. W. H. Wagner, Jr., Department of Botany, U ity of Michigan, Ann Arbor, Michigan 48104, is Librarian and Curator. ibabes — books and specimens at any time , the borrower paying all shipping costs. Mr. Neill D. Hall, arbre yp “or nga Seattle, Washington 98115, is Director. Spores exchanged and cohesion tots oat ok eae Gifts and Gifts and bequests to ee ae Sages Sos spend Se eerviens #5 eens bers and to others interested in erns. Herbarium specimens, > weloomed, and back issues ofthe Jounwar, and eaah or other pagent tax-deductible. Inquiries should be tr akimd isms American Fern Journal Vou. 60 JANUARY—Marcu, 1970 No. 1 Aberrant Leaves on Angle-Shoots of Selaginella martensii Spring! Terry R. WEBSTER Dorsal and ventral angle-meristems, which normally develop into roots (Webster and Steeves, 1967), are present at most points of shoot branching in the heterophyllous species Selaginella martensit Spring. But in stem segments that are surgically isolated from the intact plant and which are incubated on moist filter paper, these angle-meristems may develop as angle-shoots (Williams, 1937; Webster, 1969) (Fig. 6). In a recent study of angle-meristem development in excised stem segments of S. martensit (Webster, 1969), most leaves (microphylls) on angle- shoots were normal, according to the description by Harvey- Gibson (1897). But some, particularly those near the base of angle- shoots, were abnormal. The morphology and possible significance of these aberrant leaves is presented in this paper. Normally, the larger, ventral leaves of the angle-shoots are lanceolate, with a narrow, bluntly pointed apex (Fy. 1). The smaller, dorsal ones are ovoid oblique and end in a long, stiff awn (Fig. 2). Both dorsal and ventral leaves possess marginal teeth and are asymmetrical at the base. A single median vascular strand is present, and a ligule occurs at the leaf base on the-adaxial surface. Conspicuous rows of stomata occur along the vascular strand (Fig. 7). 1 This research was supported by National Science Foundation Grant GB-7068 and by a grant from The University of Connecticut Research Foundation. Volume 59, No. 4, of the JouRNAL, pp. 137~176, was issued Jan. 14, 1970. AMERICAN FERN JOURNAL VouuME 60, PLATE 1 3 CLEARED AND STAINED LEAVES OF SELAGINELLA MARTENSII NGLE-SHOOTS SHOWING VASCULAR STRANDS 5 ABERRANT SELAGINELLA LEAVES 3 In contrast to most leaves on angle-shoots, the first few leaves toward the base, proximal to the first branching of the angle- shoot, are aberrant (Figs. 3, 4, and 6). In addition to these abnormal basal leaves, a few cases of unusual leaves were found more distally on angle-shoots. Both dorsal and ventral aberrant leaves vary in shape. Instead of being entire, the leaves are variously lobed or divided at the tip (Figs. 3 and 4). In some cases they are deeply cleft. Unlike normal leaves, the base of aberrant leaves is often symmetrical. For anatomical study, angle-shoots were cleared in 2% sodium hydroxide and were stained with basic fuchsin according to the method described by Jacobs (1952). Instead of the normal single vascular strand, several strands occur in aberrant leaves (Figs. 3 and 4). As many as seven strands were observed in a single leaf. In most leaves one strand could be traced from the stem vascular system toward the leaf base. Near the base of the leaf the strand is divided, usually dichotomously, one to several times. A few ventral leaves were observed in which two separate strands could be traced from the stem into the leaf base. In neither normal nor aberrant leaves is the departure of leaf traces associated with a gap in the stem vascular cylinder. In normal and in most aberrant leaves the vascular system is closely associated with a single ligule. However, in some deeply cleft leaves, two separate ligules occur (Fig. 5) and one or more vascular strands are associated with each ligule. Ball (1925) reported sporophylls of S. martensit which possessed two ligules, one of normal size and an additional smaller ligule. Double ligules found in the present study were of normal size (F2g. 5). In aberrant leaves each vascular strand has stomata associated with it, and, compared to normal leaves, more of the leaf surface is covered by stomata (Figs. 7 and 8). The arrangement of stomata in distinct rows is not so evident as in normal leaves. Fic. 1. NoRMAL VENTRAL LEAF, X 23. Fic. 2. NORMAL DORSAL LEAF, X 22. Fic. 3. ABERRANT VENTRAL LEAF, X 46. Fic. 4. ABERRANT DORSAL LEAF, X 42. Fig. 5. ABERRANT DORSAL LEAF WITH TWO LIGULES. SOME STEM TISSUE VISIBLE THROUGH LEAF, X 47. AMERICAN FERN JOURNAL VotumeE 60, PLatEe 2 Prats 2. Fig. 6. DIAGRAM OF VENTRAL VIEW OF STEM SEGMENT BEARING AN ANGLE-SHOOT. NOTE SINGLE ABERRANT VENTRAL LEAF WITH- BRANCHED VASCULATURE TOWARD BASE OF ANGLE-SHOOT. FiG. 7. STOMATA ON NORMAL LEAF FROM MAIN SHOOT. Fic. 8. STOMATA ON ABERRANT LEAF FROM ANGLE- sHooT. The abbreviations are: L = tiGuLe. § = STOMA, V = VASCULAR STRAND. ABERRANT SELAGINELLA LEAVES 5 DISCUSSION Grambast and Rosello (1965) described leaves of S. willdenovit (Desv.) Baker in which there were two or three veins from the dichotomous branching of a single vascular trace. According to their illustrations, the leaves themselves were not so distinctly lobed or divided as the aberrant leaves described here in S. martenstt. The authors made no mention of stomata or ligules in the leaves of S. willdenovii, but they did discuss possible phylo- genetic implications of the branched vasculature. They cited fossil lycopods (Protolepidodendron Krejci, Duisbergia Kraus. and Weyl., and Sigillaria scutellata Brongn.) which possessed leaves with dichotomously branched vascular strands, and sug- gested that in S. willdenovit microphylls with dichotomizing veins represent a primitive condition. According to their view, micro- phylls with a single vein could have been derived by reduction. They suggested that, like megaphylls, microphylls may, in some cases, have been derived from a system of axes. One could apply a similar interpretation to the microphylls of S. martensii. Aberrant leaves with branched vasculature could be interpreted as representing a primitive condition, and normal leaves with a single vascular strand a more advanced condition. A comparison of normal and aberrant leaves would suggest that during evolution lobing, number of vascular strands, and rows of stomata were reduc According to the theory of recapitulation (ontogeny, in abbreviated fashion, repeats phylogeny), the position of leaves on angle-shoots could be used to indicate a primitive or an advanced condition. Although usually applied to progressive development from juvenile to adult leaves in sporelings or seedlings, Takhtajan (1959, p. 91) has applied this theory to leaves occurring on acces- sory shoots. Angle-shoots would fit into this latter category. According to the theory, the progression of microphylls in angle- shoots represents at least some stages in the evolution of micro- phylls for S. martensit. Thus aberrant leaves occurring at the base of angle-shoots would represent a primitive condition, and more distal (normal) leaves an advanced condition (Fig. 6). However, 6 AMERICAN FERN JOURNAL as Foster and Gifford (1959, p. 9) have stated, whether juvenile leaves provide reliable evidence to the morphology of ancestral leaf types is questionable, and hypotheses based on the theory of recapitulation should always be made with great caution. It should also be noted that the features found in aberrant leaves of S. martensii are not known in fossil representatives of Selaginella or in Selaginellites. At the present time there is no strong evidence to indicate that aberrant microphylls in Selaginella do in fact represent a primitive condition. Rather than having any phylo- genetic significance, the aberrant leaves in S. martensiz merely may be the result of abnormal development during the early ontogeny of angle-shoots. LITERATURE CITED Ba, N. G. 1925. On the occurrence of a double ligule in Selaginella martensii Ann. Bot. 39: Foster, A. 8. and Girrorp, E. M. 1959. nie panne Morphology of Vascular Plants. W. H. Freeman, San Franc Grampgast, L. and S. Rose..o. 1965. Les seen ons de microphylle et de méga- phylle et leur signification phylogénétique. Compt. Rend. Acad. Sci., Paris 261: 5183-5186. ee R. J. 1897. Contributions towards a knowledge of ‘66 tomy of the genus Selaginella Spr. III. The leaf. Ann. Bot ot "| Jacoss, W. P. 1952. The role of auxin in differentiation of xylem around a wound. Amer. J. Bot. 39 : 301-309. TakuTasan, A. L. 1959. Essays on the evolutionary morphology of plants. , (Transl. ie H. Gankin) Amer. Inst. Biol. Sci., Washington, D.C. Wesster, T. R. 1969. An investigation of angle—m priatoin development in excised sa segments of Selaginella martensii. Canad. J. Bot. 47: 717-722 A. Sreeves. 1967. Developmental morphology of the root of Selaginella martensii Spring. Canad. J. Bot. 45: 395-4 Wiuuiams, S. 1937. — phenomena and hormones in Gclaginelia. Nature 139: SYSTEMATIC AND ENVIRONMENTAL BroLoGy, UNIversITY OF Connecticut, Storrs, Conn. 06268. TYPE OF PLATYCERIUM ALCICORNE 7 A Further Note on the Type of Platycerium alcicorne C. V. Morton Several years:ago I published a paper, “The Nomenclature of a Madagascarian Platyceriyfn” (Morton, 1964), in which I con- cluded that A crostichum: ‘alcicorne Swartz (Syn. Fil. 12, 196. 1806) was an illegitimate change of name for Acrostichum stemaria Palisot (Fl. d’Oware et Bénin 3: ¢. 2. 1805), which was cited by Swartz as a synonym. This was an error on my part, occasioned by the fact that nowhere in the ‘Index Filicum” or in Willdenow or Desvaux is there any indication that Swartz had published his Acrostichum alcicorne prior to 1806. Mr. De Joncheere was the first to note that the name A. alcicorne Swartz was actually published in Schrader’s “Journal fiir die Botanik,” (1800, vol. 2, part 1, p. 11. 1802).1 Since this 1802 publication is prior to the publication of A. stemaria Palisot it is clear that A. alcicorne Swartz was not a renaming of A. stemaria, and must therefore have a different type. Swartz’ 1802 paper “Genera et Species Filicum” is a sort of preliminary to his extensive “Synopsis Filicum”’ of 1806; it is very brief throughout, with short diagnoses and no citation of speci- mens, localities, or synonymy in general. In the case of A. alcicorne Swartz did cite ‘““Neuroplatyceron Pluk. am. phyt. t. 429, f. 2.”” De Joncheere (1967) has argued that this Plukenet plate must be taken as the lectotype of A. alcicorne. However, it was not Swartz’ habit to give names indiscriminately, if ever, to plants known to him only from pre-Linnaean drawings. In this case there is ample evidence that the protologue included not only this pre-Linnaean plate of Plukenet but also herbarium material, for Swartz described the fertile fronds, which were not depicted by Plukenet. In cases like this there is general agreement that species ought to be typified by specimens when possible and not by plates. 1 The date “1801” on the title page has usually been accepted, but Stafleu his ‘Taxonomic a ature,’ p. 432. 1967, has shown that this part was tliched in March, 1 AMERICAN FERN JOURNAL VoLuME 60, PLATE 3 Museum botanicum Upsaliense : * liek Mieetios, LEcTOTYPE OF ACROSTICHUM ALCICORNE, NEST LEAVES (UPS) TYPE oF PLATYCERIUM ALCICORNE 9 Dr. Schelpe searched the herbarium in Stockholm and could not find a specimen suitable for a lectotype, and De Joncheere stated that Schelpe also searched the herbarium in Uppsala without finding any specimens. At my request, Dr. John Wurdack re- cently found that there are indeed specimens in Uppsala, and through the courtesy of Dr. Rolf Santesson I now have these on loan. The specimens mentioned are in the original herbarium of Afzelius. They are fine specimens, including nest-leaves and fertile leaves, mounted on five sheets. They are indicated as being from Sierra Leone collected by Smeathman. In his 1806 publication Swartz mentions (p. 196) collections from Madagascar, Commerson; Oware, Palisot; Java, Thunberg; Guinea, Afzelius; and Nova Hollandia, Née,” and on p. 12 cites ‘‘Sierra Leone” as one of the localities. In this 1806 publication Swartz had added as syno- nyms A. stemaria Palisot and A. bifurcatum Cav., and it is likely that the collections of Commerson and Palisot were cited from Palisot rather than actually seen by Swartz and that the Née cullection was cited from Cavanilles. No specimen from Java collected by Thunberg has been found. The Afzelius collection from Sierra Leone mentioned above is the only herbarium speci- men that can definitely be associated with a collection cited by Swartz. The name alcicorne is in the hand of Swartz. The five sheets in the Uppsala Herbarium are here designated as the lectotype of A. alcicorne Swartz. According to Schelpe, there is a specimen in Stockholm from Sierra Leone annotated by Wikstrém as A. alcicorne, and this is doubtless an isolectotype, although Afzelius is not definitely indicated as the source. The specimens from Sierra Leone designated as lectotype are identical with A. stemaria Palisot from Oware [Nigeria], the type of which I have seen in Paris. The Plukenet plate was apparently drawn from a specimen now in the Sloane Herbarium in the British Museum, and according to De Joncheere came from the Comoro Islands and is identical with the plant described as Acrostichum alcicorne Willemet, now properly Platyceritwm vasset Poisson. The lectotype chosen above agrees well enough with Swartz’ AMERICAN FERN JOURNAL VouuME 60, PLATE 4 Museum botanicum Upsaliense Z ee, eed Le com ss ' b gs bee ees ve. P00 Ee Ja & LecTOTYPE OF ACROSTICHUM ALCICORNE, FERTILE FROND (UPS) TYPE OF PLATYCERIUM ALCICORNE ll brief diagnosis, at least as well as that diagnosis agrees with P. vasset. The present lectotypification is really best for stability as well as fully justified by the facts. Although A. alcicorne Swartz was prior to A. stemaria Palisot by three years, it was itself illegiti- mate, being a later homonym of A. alcicorne Willemet (1796), a species based on an entirely different type (Madagascar, Stadt- mann). The next earliest name for the West African species is thus A. stemaria Palisot, and the correct name for it is Platycerium stemaria (Palisot) Desv., of which P. alcicorne (Swartz) Desy. is thus a taxonomic synonym. This is a fortunate turn of events, since P. slemaria is a name well known among horticulturists. Incidentally, Dr. Schelpe has suggested in a letter to Barbara Joe that the epithet “‘stemaria” ought to be corrected to “stemmaria,” since the root is from a garland, in Greek. This may well be, but the correction of the original spelling of a name ought not to be made for purely etymological reasons if there is no reason to suspect a purely typographical error; there are many poorly or wrongly formed names that must nevertheless be accepted. The name P. alcicorne Desv. adopted by De Joncheere for the East African and Madagascarian species is thus incorrect, which is fortunate, since this epithet has been widely misapplied to three different species—the West African P. stemaria, and the East African and Madagascarian P. vassei, and the Australian P. bifurcatum. Another species of Platycertum that may be mentioned here incidentally is P. angolense Welw. ex Baker in Hook & Bak. Syn. Fil. 425. 1868. The name appears as a comment under P. aethiopi- cum Hook. as follows: ‘‘Dr. Welwitsch has furnished us with a plant (P. angolense Welw. MSS.) which is most like this, but has a broad-cuneate fertile frond 9 in. br. at the top, without either forks or horns, and with a patch of fruit nearly as broad as the lamina.” Dr. Schelpe has suggested in a letter to me that this name ought to be rejected by Art. 34 of the Code as not being accepted by the publishing author. In my opinion it is valid, as valid as a 12 AMERICAN FERN JOURNAL large number of other names that are published rather casually in footnotes or text-notes. Baker did not either definitely accept it or reject it. He certainly did not conclude that it was exactly the same as P. aethiopicum, but gave its differences as he saw them. Since there is a name and a description, the name ought to be accepted as validly published. As it turns out, Welwitsch was quite right that it was different from P. aethiopicum Hooker, and P. angolense Welw. ex Baker is the correct name for the third African species of Platycerium, the other two being Platycerium stemaria and P. vasset; a taxonomic synonym of P. angolense is P. elephantotis Schweinf. and another probable synonym is P. velutinum C. Chr., as discussed by Barbara Joe (1964) in her useful review of Platyceriwm. LITERATURE CITED JoE, BaRBARA. 1964. A review of the species of Platycerium (Polypodiaceae). Baileya 12: 69-126 JONCHEERE, G. J. DE. 1967. Notes on Platycerium Desv. I. Nomenclature and typification of the genus and species in Desvaux’s original publica- tion of 1827. Blumea 15: 441-4 Morton, C. V. 1964. The nomenclature of a serrata Platycerium. ‘Baileys 12: 36-38. National Museum or Naturat History, WASHINGTON, D.C. DRYOPTERIS GAMETOPHYTES 13 Gametophyte Ontogeny and Sex Expression in Dryopteris ludoviciana! MricuHak. I. Cousens AND Harry T. Horner, Jr. Although the ontogeny of fern gametophytes has a long history of morphological investigation, sex expression in ferns has only recently been studied by experimental morphologists and geneti- cists. Most homosporous ferns have bisexual gametophytes whose mitotically derived gametes produce a completely homozygous sporophyte after selfing (Klekowski and Baker, 1966). Wilkie’s work (1956), which demonstrated self incompatibility in Pteridiwm aquilinum, has stimulated the use of fern gametophytes in genetic research. Klekowski and his associates (1968, 1969) used genetic and morphological criteria to characterize pteridophyte gameto- phytes in terms of their probability for intragametophytic (func- tionally bisexual) and intergametophytic (functionally unisexual) mating. This study was undertaken to determine whether the ontogeny of Dryopteris ludoviciana (Kunze) Small gametophytes differed from that reported for other dryopterid and thelypterid gameto- phytes (Kny, 1895; Waldemann, 1928; Nayar and Chandra, 1963), and to determine sex expression under controlled conditions. The effect of population density on sex expression was tested by culturing single isolated gametophytes and gametophytes in moder- ate and dense populations. The ontogeny and sex expression of Thelypteris dentata (Forsk.) E. St. John was also observed for comparison. Metuops and MATERIALS Dryopteris ludoviciana occurs in cypress and gum swamps, as well as in lime sinks, from Florida west along the Gulf Coast to Louisiana and north as far as North Carolina (Brown and Correll, 1 This p r is a portion of a thesis completed by the senior author for the Master of git Degree, I. S. U., 1969. 14 AMERICAN FERN JOURNAL 1942). A detailed record of its occurrence is not available, but it is less common than other Dryopteris species. Two Dryopteris ludoviciana sporophytes were greenhouse- grown after their collection on November 20, 1966, in Alachua County, Florida, in the Devil’s Millhopper, a lime sink 5-6 miles northwest of downtown Gainesville. A voucher specimen (John T. Mickel 1730) is in the Iowa State University Herbarium. The plants used in this study were diploid (2n = 82) (A. R. Smith, pers. comm.). Fertile pinnae were cut from vigorous fronds and rinsed in running distilled water to remove foreign spores. The pinnae were then sealed in small manila envelopes. After most of the sporangia had dehisced (8-12 days at 20°C) the spores were trans- ferred to small corked vials and utilized within two months. All spores were sown on a sterile mixture of loam and coarse sand (4:1). Varying numbers of spores were sown to produce different densities. Population density was determined by counting all gametophytes visible at 30. The average number of gameto- phytes per cm? was then determined. Dense populations contained 10-21 gametophytes per em? and moderate populations had 1-3 per cm’. Isolated spores were cultured in small petri dishes. Culture vessels containing only sterile soil were opened for watering and observation with the same frequency as the experi- mental cultures. Fern gametophytes appearing in these cultures reflected the extent of contamination (about one per cent). Densely sown cultures of Thelypteris dentata were also maintained under the same conditions of culture as Dryopteris ludoviciana and later were surveyed for the occurrence of gametangia. Conditions of culture, including the use of soil as a substrate (Atkinson and Stokey, 1964), were selected to approximate the habitat of naturally occurring fern gametophytes. The cultures were maintained in a Percival growth chamber. Diffuse illumina- tion was provided by white fluorescent tubes and incandescent bulbs at an intensity of 350-400 ft-c. Temperature at the soil surface was maintained at zak during the 12 hr light and 12 hr dark periods. DRYOPTERIS GAMETOPHYTES 15 Samples of 24 or more gametophytes were removed for observation of development and sex expression at 25, 35, 45, 58, 72, and 111 days after sowing. An effort was made to select gameto- phytes representative of all sizes and stages. The gametophytes were placed in a petri dish or on a glass slide containing 1.75% lactose solution for microscopic examination. This medium pre- vented plasmolysis for 8-12 hours and did not hinder the dehis- cence of antheridia or the motility of sperms. GAMETOPHYTE ONTOGENY AND GAMETANGIA Spores of Dryopteris ludoviciana are bilateral with a prominent perispore. A spring-collected sample (April to June) of 162 spores ranged from 24-50 yu long by 16-33 » wide. One hundred and eight fall-collected spores (October) ranged from 21-54 u long by 16-37 uw wide. The spring-collected spores averaged 36 X 24 uw and the fall-collected spores averaged 36 X 26 yw. Spore size was con- siderably less uniform in the fall collection (Fig. 1), but the bio- logical significance of this, if any, is not known. A bimodal dis- tribution of spore size was not found. Most spores germinated within 6-20 days, but some took as long as three months. The spore coat ruptured irregularly or along the ridge and remained attached to the basal cell (Fig. 2). The first rhizoid was delimited shortly after germination, and a second rhizoid sometimes formed before filamentous growth began (Fig. 3). Successive divisions then produced a 4—6-celled filament which was either stout (Fig. 4) or elongate (Fig. 5). Filamentous growth was prolonged if a spore germinated in a soil crevice, or was otherwise shaded (Fig. 7). These attenuated filaments eventually became planate. Longitudinal or oblique divisions in the terminal cell of the filament initiated planate growth (Fig. 6). These divisions could occur when the filament was only two or three cells long (Fig. 8). Subsequent longitudinal divisions could occur in all but the basal cell of the filament, obscuring the earlier filamentous form (Fig. 9). Continued divisions in two dimensions were most frequent distally, producing a triangular gametophyte of 10-15 cells with 16 AMERICAN FERN JOURNAL marginal trichomes (Figs. 11, 12). This stage was occasionally irregular (Fig. 10), but these irregularities were obscured by sub- sequent symmetrical growth. The fastest growing gametophytes established an apical cell 12-16 days after germination (F7g. 13; MNZODO-E Z— THO—-=EF MBIOVYH N @ 20 22 24 26 28 30 32 34 36 38 40 42 44 46 48 50 52 54 SPORE LENGTH IN MICRONS Fig. 1. Size DisTRIBUTION OF DRYOPTERIS LUDOVICIANA SPORES. © = ONE SPRING-COLLECTED SPORE; @ = ONE FALL-COLLECTED SPORE. Table I). Slower growing gametophytes reached this stage within 16-30 days. The apical cell cut off derivatives laterally to produce the notch meristem 14-32 days after germination (Figs. 14, 15; Table I). Once this stage was reached, irregularities in growth were rare, and all gametophytes, except ameristic males which lack a notch meristem, became cordate (Fig. 16). Trichomes were common on the dorsal surface, averaging one trichome per five prothallial cells. This ratio diminished to one DRYOPTERIS GAMETOPHYTES 17 trichome per fifteen prothallial cells on the wings and to one per 20-30 prothallial cells on the ventral surface. Gametophytes 20-45 days old and 0.5-0.7 mm wide initiated a cushion which was two or three cells thick at first and later four or five cells thick. Lateral prothallial cells (Fig. 20) became much larger than younger anterior cells (Fig. 19). Older gametophytes were 0.9-13.0 mm wide (Figs. 17, 21), depending on their pattern of sex expression. TaBLeE [. INITIATION (IN DAYS AFTER GERMINATION) OF THE STAGES OF DEVELOPMENT OF DRYOPTERIS LUDOVICIANA GAMETOPHYTES. Rapidly growing Slow growing Developmental stage gametophyte gametophyte Filamentous 6 6 First longitudinal division £¢ 7 Early planate (9-15 cells) 8-11 11-29 Apical c ell 12-16 17-30 Notch meristem 14-19 20-32 Cushion initiation 20-26 27-45 The antheridia are of the polypodioid type (Atkinson and Stokey, 1964), consisting of a single basal cell, a cylindrical middle cell which surrounds the central primary androgonial cell, and an undivided cap cell. The archegonia are of the advanced type (Atkinson and Stokey, 1964) with four or five tiers of neck cells, each tier con- taining four cells. The central canal cell was binucleate and en- larged distally. The archegonium was slightly larger at its apex than at its base and was always inclined slightly toward the base of the gametophyte. Archegonia occurred ventrally on a broad area just behind the notch, and up to 80 occurred on gameto- phytes 8-13 mm wide. Sex Expression IN POPULATIONS Dryopteris ludoviciana has a normal sexual alternation of generations. The gametophytes observed in isolated, moderate, AMERICAN FERN JOURNAL VoLuME 60, PLATE 5 Dr OX og At AY Puate 5. Earty OntoGeny or D. Lupoviciana GAMETOPHYTES. THE NUMBERS IN CIRCLES INDICATE DAYS AFTER GERMINATION. THE MOST COMMON SEQUENCE OF DEVELOPMENT Is Fias. 2, 4, 6, 11-15. OnE cm = 100y FOR Figs. 7, 15; 1.5 cm = 100 uw FoR ALL OTHER FIGURES. DrYOPTERIS GAMETOPHYTES 19 and dense cultures expressed one of five possible patterns of sexuality, or lacked gametangia. The five sex expression patterns were based on the presence of one or both types of gametangia, and, where both kinds of gametangia occurred, the sequence of their appearance. The presence or absence of a notch meristem was also recorded for all gametophytes. Rapidly growing gametophytes (Table I) usually became uni- sexual females and were the first plants in all populations to become sexual (Fig. 18). Archegonial initials were found on 24-day-old gametophytes as small as 0.9 mm wide. These gametophytes continued to grow and produce additional archegonia, attaining an average width of 4.6 mm 45 days after germination (Fig. 21). These gametophytes remained unisexual after 105 days and became 8-13 mm wide (F7g. 17). Older gametophytes occasionally produced basal outgrowths which became antheridial. Protandrous bisexual gametophytes are typical of most homo- sporous leptosporangiate ferns (Atkinson and Stokey, 1964), but were rare in D. ludoviciana populations studied here. These gametophytes first produced antheridia among the rhizoids and on the basal half of the cushion. Six to 20 days later archegonia formed below the notch meristem. Antheridia numbered fewer than 80 on protandrous bisexual gametophytes. Twenty of the 527 ceramic surveyed were iid eaten 9 bisexual. In bz. sexual archegonia occurred in the VeVitia unite typical position below the notch meristem and antheridia were restricted to the distal half of the cushion adjacent to and among the archegonia. Maturation of both kinds of gametangia was synchronous, so that the periods of antheridial dehiscence and archegonial receptivity overlapped. These gametophytes first became sexual 30-40 days after germination and continued to produce both kinds of gametangia. After 45 days a gametophyte bore an average of ten archegonia and 22 antheridia. Ameristic male gametophytes developed antheridia precociously but failed to develop an organizing notch meristem. They became sexual shortly after germination, producing antheridia on almost every cell, and occasionally two per cell. Ameristic males devel- AMERICAN FERN JOURNAL VouumME 60, PLATE 6 as spegute'ctt Gant ces an wen \ 1 On? a THAR 8 POG: . 1 mm Piate 6. Late Ontogeny or D. LupovicianA GAMETOPHYTES. THE NUMBERS IN CIRCLES INDICATE DAYS AFTER GERMINATION. Fic. 16. SLOW GROWING GAMETOPHYTE WITH CUSHION BUT WITHOUT GAMETANGIA. Fic. 17. PHYTE WITH ARCHEGONIA. Fic. 19. CELLS FROM ANTERIOR MARGIN OF WING; cF. Fig. 20. Fie. 20. CELLS FROM LATERAL AREA OF Fia, 21. Fic. 21. RAPIDLY GROWING UNISEXUAL FEMALE GAMETOPHYTE. DRYOPTERIS GAMETOPHYTES 21 oped only from spores which either remained within sporangia, were next to sporangial fragments, or were in contact with the rhizoids of older gametophytes. Their form varied from filamentous to spathulate. SPRING FALL SEXUALITY fed pa pe | eed oe Loe 5S 35 25 35 45 an 8S: | 45 23 35 «645 25 35 45 23.6 633 «C45 i] . FEMALE 28 - 75134 - 691.9 - 2.69 43 60 80] 13 18 20 | 1.6 3.3 6.1 o PROTANDROUS BISEXUAL 0 - 0 oO = 8 - - - 0 0 0 o 0 0 - = BL Jf S¥8cHRonous BrsexvaL hs 2 = PES Ss ae 02 ato 7 2 - 3.96.3 A AMERISTIC MALE 0 - 0 Go = 6 - - - 0 0 4) 000 - - - TV § MERISTIC MALE 0 - oO G'= 6 a ae o 0 Oo 000 at. ice re E GAMETANGIA ABSENT 71 - 13 90 - 1 0.4 - 4.0 oS & aes i 5 Ly 33, 3 0.4 0.5 0.8 D TOTALS 100 -100]126 - 8] - - - [ff 100 100 100 | 30 30 25 a ME eae pba = senile ae ete a ee 34 40 399 10 14 11 1.52.9 3:7 o PROTANDROUS BISEXUAL - - - - - - - - - 0 0 - O- - 7 - - “ SINCKAONDS BIMKUAL B=, = Sy AS nee = 7 28 3%] 21010 | 1.8 2.5 3.1 R AMERISTIC MALE - we Or - - - - - 0 0 o 0- 0 - - * A MERISTIC MALE 7 - - ~ - - - - 7 11 = 0 3.2 9 0.61.4 - T Bf caerancia ABSENT ae oe eh ees ee 48 30 2571511 7 | 0.5 0.3 0.9 E TOTALS ~~ <=» fF. «=9"%~ = ~ F100 100 100 | 30 36 28 mee aah FEMALE 43 = 30 21 - 2295.3 - 3.68 40 30 32 | 29 27 26 | 2.4 3.0 3.5 1D |] PROTANDROUS BISEXUAL eet S| a eae ket ae Os £4 6 ot = = 320 EJ syvcnronous BIsexuaL ee ome es S..4S) 22.0 6 01 29 3 N IC MALE ico ke P-2 = PO OE OO: 680 ht ieee $F cristic me 33 - 20716 - 1491.7 - 1.5 | 20 28 36 | 14 25 29 | 0.8 0.6 0.9 E GAMETANGIA ABSENT 10° = 368 oS 96 2.0} 2.8 I oe et 8 pase 7 | 0.6 6:8 0,5 TOTALS 100 - 100149 - 73] - - - [100 100 100 | 72 89 81 fa Og TaBLe IT, Sex Expression In 25,35, AND 45-DAY-OLD IsoLATED, MODERATE, AND DENSE DRYOPTERIS LUDOVICIANA CULTURES. Meristic males produced 100-300 antheridia after they became cordate. The antheridia occurred on two-thirds of the wings, and 22 AMERICAN FERN JOURNAL extended to within four cells of the notch meristem. Meristic males continued to produce antheridia after being transplanted back to soil, and were not bisexual after 105 days. omMarPron~— 4 il Mis 25 Oars 35 Days 45 bars M 0 D . E R A T E © TUT $680 ao wo S480 of no ve PUT $64 6 6 wo 2S Days {he @ m0 aS oars 1000-2500 m i 5 8 1 4 igs 47 Above 2500 m 0 3 5 2 3 15 54 Abaxial Surface 1000 m () 0 3 0 1 0 26> 1000-2500 m 1 4 18 Zz 5 7 19 Above 2500 m 0 2 ae 4 5 3 10 *See Table IT for scale types and references to Located ag: in Bell, 1955. » Based on 13 species, 4 of which were analyzed as to type. surfaces at higher elevations. However, this trend appears to be reversed on the adaxial surface. Paraphyses.—Copeland (1947, p. 119) stated that paraphyses are wanting in Elaphoglossum. However, recent studies by Ander- _ son and Crosby (1966) indicate the presence of paraphyses in some Hawaiian species. A survey of fertile fronds in Costa Rican material indicates that 56% have paraphyses. In most cases the paraphyses are short, uniseriate, multicellular branches arising from the upper portion of the sporangial stalk near the base of the sporangium. In a few cases they arise from the receptacle MorPHOLOGY OF ELAPHOGLOSSUM 79 (Lloyd 4268) or from both the receptacle and the sporangial stalk (Lloyd 4284). They are usually glandular at maturity. Paraphyses occur in 68% of epiphytic species and only 43% of terrestrial (Table IT). TaB_e lV. PERCENT OF SPECIES WITH CHARACTERISTICS IN RELATION TO ELEVATION Character 0-1000 m 1000-2500 m 2500-above Swelling Present 15 84 68 Jointed 75 90 79 Pneumatophore: As 25 a 11 B 0 13 4 C 0 45 36 D 0 8 7 E 25 a fe F 25 13 7 G 25 11 28 Paraphyses Present 100 54 61 No. species investigated 4 38 28 «See Table II for description of types. » Based on percent of taxa with fertile fronds. Asexual Reproduction. Vegetative reproduction is infrequent in most ferns and rare in Elaphoglossum. In the Costa Rican material it was found in only one species, E. undulatum (Willd.) Moore (Lloyd 4177). Plants of E. undulatum occur in very wet secondary forest near San Ramon and regularly produce asexual buds from the terminal portion of the midvein of the frond. This species was usually epiphytic, normally between 0.5 and 1 meter above ground level, with pendent fronds which frequently came in contact with the soil. Buds and new plantlets were observed on mature fronds, increasing in size as the parent fronds aged. As the parent fronds degenerate, the new plantlets formed ter- restrial colonies at the base of the tree. SO AMERICAN FERN JOURNAL Other Sporangial Features.—Sporangia tend to arise in numerous, circular sori below the veins. As the fertile fronds mature, these sori merge together and become acrostichoid. The number of indurated annular cells of the sporangia vary from 10 to 15, with 12 being the most frequent. Chlorophyll is present in the sporangial stalk in about one-third of the species. Discussion The most obvious trend found in this study in Elaphoglossum is toward the terrestrial habitat with increasing elevation. In lowland rain forest, 93 % of the species were epiphytic. In highland wet forest (above 2500 m) only 50% of the species were epiphytic. This trend is frequent in other groups of plants, including Bryo- phyta, Bromeliaceae, and Orchidaceae. At lower elevations species are either terrestrial or epiphytic, but not both. At upper eleva- tions, particularly in exposed habitats, this distinction breaks down. The cause of this phenomenon is only vaguely known. Wet cloud forest, with its somewhat stunted trees, has altered the terrestrial habitat and permits normally epiphytic species to grow. This may be due to increased humus, mosses, water and light. At lower elevations the available moisture in the soil is greater than that on the trees, but here other factors, such as lack of light or available substrate, must control plant distribution. There is little evidence to indicate whether the terrestrial habitat is less extreme than the epiphytic, although it is apparent that the former may provide the more suitable habitat during dry periods in those areas with a marked wet-dry climate. Terrestrial habitats also provide greater protection from drying winds. Some terrestrial habitats, especially those of disturbed areas along roadsides where there is no protective cover, appear to have more extreme condi- tions. Species of Elaphoglossum that occur in these habitats have coriaceous fronds, a feature that Lloyd (1965) found to be as- sociated with xerie environments in Polypodium. MorpuHo.ocy or ELaAPHOGLOSSUM 81 There are certain characters correlated with either epiphytic or terrestrial habitat. These are: Epiphytic Terrestrial Non-jointed Jointed Non-swelling Swelling Type C pneumatophore Type G pneumatophore Pneumatophores present Pneumatophores absent Scaly Glabrous Type B, C, & E scales Type C scales (adaxial surface) (adaxial surface) Paraphysate Non-paraphysate The strongest of these trends is toward the absence of para- physes in terrestrial plants. In addition, there is a slight increase in the frequency of paraphyses with higher elevations, although the number of terrestrial species also increases. If paraphyses evolved as protective structures against desiccation of young sporangia, epiphytic plants more exposed to changing conditions would be expected to show a higher frequency of presence. This might also be true of plants in more extreme situations at higher elevations. Bell (1955) found a close association between the joint and the presence of pneumatophores, although he believed that they were probably independently controlled genetically. The function of the pneumatophore is unknown. The function of the joint may be as a mechanism for shedding leaves, as in Oleandra. Holttum (1966) believed that this was an adaptation to the epiphytic habit. However, in Elaphoglossum, terrestrial plants show a higher frequency with joints than do epiphytes. Species at higher eleva- tions also possess this structure less frequently. The trends in scaliness are of uncertain significance. Plants with both glabrous and densely scaly fronds occur side by side in many different habitats. Christ (1899) believed that species at higher elevations tended to have densely scaly fronds and those at lower elevations glabrous fronds. My results indicate that the adaxial surface of the frond is less densely scaly at higher eleva- tions than at mid-elevations. However, on the abaxial surface 82 AMERICAN FERN JOURNAL scale density increases with elevation. Over 50% of the species from elevations above 2500 m had glabrous fronds or fronds with less than ten percent of the surface covered by scales. It is obvious from this study that the relationship between the characters investigated and the habitat is complex. The wide diversity in structure may be due to maintenance of much of the diversification through time since the origin of the genus in a stable, continuously tropical habitat (Bell, 1956), and in fact, there may be no correlation with habitat for many of the features I studied. Further studies are needed, however, before definite conclusions can be reached as to the evolution of the various features and their relationship to the environment. se of Characters for Taxonomic Purposes.—Many of the fea- tures elaborated in this study appear to be useful for taxonomic delineation of species. Of particular importance are scale type, pneumatophores, presence or absence of a joint and phyllopodium, and presence and type of paraphyses. Bell (1955) found that the scale types are constant in any one species. This may be true of the other features, although large numbers of individuals of each species have not been investigated. Nearly all 70 species can be distinguished using a combination of these characters. ACKNOWLEDGEMENTS ; Much of this work was done while I was involved in a course on | the biology of tropical pteridophytes, sponsored by the Organiza- : tion for Tropical Studies. I am indebted to that organization and to the University of Costa Rica for providing funds and facilities while in Costa Rica. I am also indebted to Drs. W. H. Wagner, Jr., J. T. Mickel, D. B. Lellinger, A. M. Evans, and C. V. Morton . for their helpful criticisms and comments during the study. Speci- | men identification was done by Drs. Mickel and Lellinger. : | : : LITERATURE CITED ROUEN, Ad R. and M. R. Crossy. 1966. A revision of the Hawaiian species Elaphoglossum. Brittonia 18: 380-397. BEL, P. = 1950. Studies in the genus cc area Schott. I. Shae struc- ture in relation to habit. Ann. Bot., Lond. n.s. 14: 545- Nores AnD News 83 —————. 195la. Studies in the genus ae II. The root and bud traces. Ann. Bot., Lond. n.s. 15: —346. —————-, 1951b. Studies in the genus see er Il. aa: oes of the rhizome and frond. Ann. Bot., Lond. n.s. 15: 347-3 —————=, 1955. Studies in the genus Flapbolossunt: IV. The “at shologttl series in the genus and their phylogenetic interpretation. Ann. Bot Lond. n.s. 19: 173-199. — . 1956. Studies in the genus Elaphoglossum. IV. Part Two. Ann. Bot., Lond. n.s. 2 —88. Curist, H. 1899. } Monographie des genus Elaphoglossum. Denkschr. Schweiz. aturforsch. Ges. 3 hoe Coren, E. B. 1947. se Filicum. Ww altham, Mass Hourrum, R. E. 1966. Florae Malesianae Preciiscres XLIV. The fern genus E aphoglossum in Malaysia, with descriptions of new species. Blumea 317-326. Luioyp, et

° 4, « @, @* -.-+6 fe ah “@ Ses. --.-3 1 =~ © “Ors6* Re ets. tees. e ~~." * J00ee-~ * . * , E. B. 1947. Genera Filicum. Chronica Botanica: Waltham, Mass. Hayata, B. 1927. On the systematic importance of the stelar system, in the Filicales. Bot. Mag. Tokyo 41:697-718. Hourrum, R. E. 1954. Flora of Malaya, vol. 2. Ferns of Malaya. Gov’t. Printer, Singapore. Iwarsuki, K. 1964a. On Hypodematium Kunze. Acta Phytotax. Gebot. 21:43-54. 64b. Taxonomy of the thelypteridaceous ferns, with special aca to the species of Japan and adjacent regions. II. Circum- scription of the group. Mem. Coll. Sci. Univ. Kyoto, B, 31:1—-10. Loyat, D. 8. 1960. Some observations on the gametophyte of Hypodematium crenatum (Forsk.) Kuhn with a note on the phyletic affinities of the genus. J. Indian Bot. Soc. 39:133-139. Meura, P. N. and D. S. Loyau. 1956. Some observations on the cytology and anatomy of Hypodematium crenatum. Curr. Sci. 25:363-364. Nayar, - ~ 1962. Morphology of the spores and prothalli i some species of Polypodiaceae. Bot. Gaz. 123:223-232. : ie 70. Clasaifiestion of the homosporous ferns. Chapter II in B. K. Nayar and S. Kaur, Companion to Beddome’s Handbook to the Ferns of British India, Ceylon & Malay Peninsula. Chronica Botanica, Delhi. , and 8. Devt. 1964. Spore morphology of Indian ferns. I. Aspidia- ue. Grana Palynol. 5:80-1 , and §. Kaur. 1965. Studies in the fern genera Bolbitis and Egenolfia. I. The adult sporophytes of some species. J. Linn. Soc Bot. 59:127-140. 118 AMERICAN FERN JOURNAL and ——————. 1968. Spore germination in homosporous ferns. J: Palyno 4:1-14. —————. 1969. Prothallial development in homosporous ae Phytomorphology 19:125-135. . Baspai, and 8. Cuanpra. 1968. Contributions to the mor- phology of the leek genus Oleandra. J. Linn. Soc. Bot. 60:265-282. NATIONAL Botanic GARDENS, LucKNow, INDIA. Shorter Note PYRROSIA PRINCEPS, A FERN New To Cuttivation.—I have recently received for identification from Mr. J. W. Peterson an unidentified fern that is being cultivated in Longwood Gardens from material collected by R. G. Wilson in New Guinea and Bougainville (Peterson J-1087, collected Jan. 12, 1970). It proves to represent a species previously unknown in cultivation and very restricted in its natural distribution: Pyrrosia princeps (Mett.) Morton, comb. nov. Polypodium princeps Mett. Ann. Lugd. Bat. 2: 232. 1866. Type: New Guinea, Zippel (holotype L, Morton photograph 833). Cyclophorus princeps (Mett.) C. Chr. Ind. Fil. 200. 1905. This is a characteristic species, known to me from only three collections (aside from the type) from New Guinea: Auga River, in clumps among rocks near river, alt. 580 m., Papua, Sept.—Nov., 1933, Brass 5492; terrestrial in edge of forest on riverbanks, Biniguni Camp, Gwariu River, Milne Bay District, Papua, July 30, 1953, Brass 23703; on limestone cliffs, Biak Island, December 2, 1945, Grether & Wagner 4220. From the data on the label of the cultivated plant it appears that the species may occur also on ugainville Island, Solomon Islands. In habit this species is quite different from the commonly cultivated Pyrrosia lingua (Thunb.) Farwell, which has long- stalked fronds seattered along an elongate, creeping rhizome. The rhizome is compact and larger in P. princeps, and the fronds are fasciculate. The fronds are probably the largest in the genus Pyrrosia, reaching more than a meter in length and 10 cm. in RecENT FERN LITERATURE 119 width. The fronds are essentially exstipitate, which distinguishes them from such species as P. grandissima (Hayata) Ching. The Indian Pyrrosia costata (KK. B. Presl) Iwatsuki and the Philippine P. splendens (K. B. Presl) Ching are similar in the frond outline, but the indument is quite different, the scales being dimorphous, the layer of peltate scales being underlain by white, dissected scales with elongate frizzy arms. In P. princeps the scales of the lower surface are monomorphous, all like, with numerous, short spreading arms.—C. V. Morton, National Museum of Natural History, Washington, D.C. 20560. Recent Fern Literature A ComMMENTARY ON SomE Type SPECIMENS OF FERNS IN THE Herparium or K. B. Prest, by R. E. Holttum. Nov. Bot. Inst. Univ. Carol. Prag. 1968 :3-57. June, 1969.—The study of fern types has in some respects only begun. Original specimens were to a certain extent neglected until the twentieth century. The earliest paper that I recall specifically on fern types is that of Christensen in 1910 on some of the types of Swartz. Probably no others were published until 1936, when Weatherby wrote a most valuable paper on the American types of Desvaux, and 1937, when Christensen wrote on the types of Cavanilles and Cesati. In 1954 I began a general study of fern types, which has continued up to the present time. This is a gigantic task I shall never finish, for I estimate that there are in excess of 50,000 fern types to be located and studied. The Czech botanist K. B. Presl described many species between 1825 and 1851. His types have been in Prague ever since but they have almost never been consulted, which is unfortunate since some of his species, although rather fully described for his day, have remained dubious. Dr. Holttum has remedied this by jour- neying to Prague to locate the Malaysian types, which are among the most important, for most of Presl’s new species were based on the Philippine Island collections of Thaddeus Haenke and Hugh Cuming. Presl’s herbarium was still unmounted and just as Presl left it when Underwood visited it in 1905, but it is now mounted and filed, partly in the general herbarium of the Bo- 120 AMERICAN FERN JOURNAL tanical Institute of Charles University and partly in the National Museum, Prague. Holttum gives an interesting account of the history of the herbarium and also of the Haenke collections obtained on the Malaspina Expedition (1789-1794). Holttum’s paper gives a commentary on all of Presl’s Malaysian collections and gives his current disposition of them. The paper contains a rather large number of typographical errors, which are not at all Holttum’s fault but which are attributable to the Czech type- setters’ lack of familiarity with English and with some standard botanical practices, such as the setting of new combinations in boldface. The most significant errors I have noted are: Page 3, paragraph 1, lines 11 and 12 are reversed. Page 21, line 16: For ‘Now identified as Drynaria quercifolia (Linn.) J. Sm.”, read “Now identified as Goniopteris tetragona (Swartz) Preal. ” T obtained the correct reading from Holttum himself, who consulted his original manuscript. Page 36, line 12: For ‘‘1962,” read “11862.” The points of interest are far too numerous to mention here, but special mention should be made of the discussion of Presl’s genera Pronephrium and Proferea, the former being an earlier name for Abacopteris Fée, if that is recognized as a genus, and the latter a synonym of Cyclosorus megaphyllus (Mett.) Ching (= Thelypteris megaphylla (Mett.) K. Iwatsuki). Holttum’s in- tensive study of the ferns of this area for more than 40 years renders him uniquely qualified to discuss the taxonomy. I would like to comment only on the treatment of five species, which are mentioned below. Page 21. Polypodium plukenetii Presl is identified as probably identical with Phymatodes alternifolia (Willd.) Presl (syn. Phyma- todes nigrescens (Blume) J. Smith). This misidentification of Polypodium alternifolium Willd. with Polypodium nigrescens Blume is a mistake of Copeland. Sledge (Bull. Brit. Mus. Nat. Hist., Bot. 2: 144. 1960) has commented that a photograph of the type in the Willdenow Herbarium in Berlin shows that alternifolium Willd. is a form of Polypodium phymatodes (i.e. P. scolopendria) rather than a synonym of P. nigrescens. He says that the type ReEcENT FERN LITERATURE 121 sheet is a small but fertile frond, by which he must have been referring to Herb. Willd. 19637/1 (which zs a small, fertile frond); however, there are two other sheets in Herb. Willd., 19637/2 and 19637 /3 (all evidently a part of the same collection, India, Klezn), and these other sheets are large plants which show that P. alternt- foliwm Willd. perhaps equals Phymatodes banerjiana Pal & Pal (Amer. Fern Journ. 53: 103. 1963), a species only recently segre- gated from P. scolopendria. I pointed this out to Dr. Pal, but he prefers to keep his species banerjiana separate (I.c. 108) on some characters that I can not follow. In any case, P. alternifolium is different from P. nigrescens; Presl’s P. plukenetti should therefore be placed as a synonym of P. nigrescens and not of P. alternifolium. Incidentally, there are plants in cultivation in the United States as P. scolopendria that appear to be surely P. alternifolium, and such plants are to be expected in botanical gardens elsewhere. The characters of these species are well stated by Pal and Pal. Page 27. The new combination Pyrrosia costata (Presl) Holttum was previously made by Tagawa and Iwatsuki (Acta Phytotax. Geobot. 22: 100. May, 1967). Tagawa and Iwatsuki cite the wrong basionym Niphobolus costatus Presl ex Beddome (Ferns Brit. Ind. ad t. 120. 1866, excl. tab.), which is a later use of the epithet costatus but which is based ultimately on the same type as Apalophlebia costata Presl. All these names are based on Poly- podium costatum Wall. List no. 265. 1828, nom. nud. Page 30. Blechnopsis malaccensis Presl is referred to Blechnum indicum Burm., which should rather be B. serrulatum L. C. Richard. In the “Index Filicum, Suppl. 3” Christensen definitely equated B. indicum Burm., previously regarded as dubious, with the common and widespread B. serrulatum, but the basis for this opinion is unknown. An examination of the type of B. indicum Burm. in Geneva shows that this species is by no means the same as B. serrulatum, but is, strangely enough, Asplenium longissimum Blume, as annotated on the sheet by Mr. F. Ballard, an identifi- cation I can confirm. Fortunately the well-known name Asplentum longissimum Blume need not be renamed “A. indicum” because there exists (but only very recently!) an Asplenium indicum 122 AMERICAN FERN JOURNAL Sledge (Bull. Brit. Mus. Nat. Hist. 3: 264. 1965), an entirely different plant. Page 33. Diplazium vestitum Presl. There were two syntypes cited—Cuming 333 and 336. Holttum cites 336 as the ‘‘Type,” but this should be “Lectotype.” He stated that he did not find 333 at Prague, which is probably to be explained by Presl’s comment (Epim. Bot. 84. 1851) under his D. grammitoides, ‘A ffinis sed diversa species est D. tenerum ex insula Leyte (Cuming n. 333).”’ This indi- cates either that Cuming 333 was a mixture in Presl’s Herbarium, one sheet being D. vestitum and one an undescribed species that he called “D. tenerum,” or that Presl changed his mind about the status of 333. Since Holttum did not find 333 filed as D. vestitum, it is likely that the latter is true, and that 333 is filed as ‘‘D. tene- rum.” Holttum states that 333 is not represented at Kew; however, it 7s there, filed as D. “sylvaticum,” (Morton photograph 18620) and I have found it also in Firenze (Morton photograph 16124). A peculiar thing is that J. Smith cited 333 as from Samar, and the Kew and Firenze sheets are from Samar and also the specimen cited by Pres] as his D. vestitum, whereas the specimen mentioned by Presl as “D. tenerum” was said to be from Leyte, which was probably an error, since the same number was not assigned to Cuming collections coming from different islands even though the plants might have been considered the same species. The speci- mens of Cuming 333 that I have seen are apparently referable to D. petiolare Presl and not to D. vestitum, which is more divided and more scaly on the rhachis. Cuming 336 was fixed as lectotype of D. vestitum by Hooker (Second Cent. Ferns t. 46. 1860 and Sp. Fil. 3: 260. 1860), who referred only Cuming 336 to Asplenium vestitum (Presl) Hooker, and cited 333 under A. sylvaticum (I. ¢- 248. 1860). Page 51. In his early and valuable work on Stenochlaena (Gard. Bull. Str. Settl. 5:259. 1932), Holttum identified a rather common and distinctive Malaysian species as Stenochlaena laurifolia Presl (Epim. Bot. 164. 1851), admittedly without having seen either the type or the original description, which is quite understandable, Presl’s books being rare and his specimens in Prague very remote REcENT FERN LITERATURE 123 from Singapore, where Holttum was working at the time. Now on seeing Presl’s type (Cuming 226) he is certain that S. lawrifolia Presl is a synonym of the common S. palustris (Burm.) Bedd., and that the species that he called ‘4S. laurifolia” in 1932 needs a name. He remedies this by proposing ‘“‘Stenochlaena cumingti Holttum, nom. nov.—S. laurifolia (non Presl) Holttum, Gard. Bull. Str. Settl. 5: (1932) 259; Copel., Fern Fl. Philip. 428.” But this is an entirely inadmissible procedure, for one can give a new name only to a validly published species that is in need of a new name; a misidentification is not a validly published species. This is not a new name but a new species, and as such it requires a Latin diagnosis and a designation of a type, neither of which Holttum provides. Therefore, S. cumingii Holtt. is a nomen nudum, not validly published. Dr. Holttum should provide a proper descrip- tion and a designation of a type to validate this name, which is needed.—C.V.M Reviews or TropicaL AFRICAN PrertpopHyTa, 1, by E. A. C. L. E. Schelpe. Contributions from the Bolus Herbarium, No. 1, published by The Bolus Herbarium, University of Cape Town, Rondebosch C. P., Republic of South Africa, pp. 1-132. 1969. Price 2 Rands.—There are at present no general treatments of the ferns of tropical Africa, although there are some fine regional treatments, such as Madame Tardieu’s “Les Pteridophytes de l'Afrique Intertropical Francaise,” (1953), pteridophytes in the “Flore du Gabon,” (No. 8, 1964), and ‘Flore de Cameroun,” (No. 3, 1964), and Alston’s ‘‘The Ferns and Fern-Allies of West Tropical Africa,” (Suppl. ed. 2, The Flora of West Tropical Africa, 1959). East tropical Africa has been unworked, except for the papers on Ethiopia by Pichi-Sermolli, and so Dr. Schelpe’s new treatment will be of great value to students everywhere. This rst part discusses the families Grammitidaceae, Azollaceae, Salviniaceae, Vittariaceae, Lomariopsidaceae, Adiantaceae, and Polypodiaceae (sens. restr.). The Grammitidaceae are considered to contain two genera, Grammitis and X iphopteris (incl. Cteno- pteris); my union of these into the single genus is termed by Schelpe ‘“‘a rather extreme standpoint,” although Schelpe separates 124 AMERICAN FERN JOURNAL these genera only by the character ‘Frond simple, entire to shallowly crenate” as opposed to ‘‘Frond pinnatifid to deeply pinnatifid,” and adduces no other characters; there are in fact no other characters. Since I do not know of any other fern genera that are separated only by the character of blade division, and that only a difference between blades shallowly crenate as opposed to pinnatifid, my viewpoint does not seem unduly extreme. In fact, many fern genera that have blades normally pinnatifid to com- pound have some species with simple blades, e.g. Adiantum, Trichomanes, Lindsaea, Polystichum, Thelypteris, Asplenium, Polypodium, in fact many large genera. The new combination Xiphopteris albobrunnea (Baker) Schelpe is proposed, based on Polypodium albobrunneum Baker (1877), but Polypodium sechellarum Baker (1874) is prior. Schelpe rejects the latter on the ground that it was based on two elements, one of the syntypes representing X. albobrunnea and the other X. villo- sissima subsp. subpinnata. Schelpe also states that the original description is not conclusive as to which element should bear the name sechellarum, and so rejects the name for either. This is contrary to the Code and to usual taxonomic procedures, which provide the workers should select lectotypes where these have not been provided. These lectotypes can be and frequently are quite arbitrary, where there is no special reason to select one rather than another element. But here there is a clear choice, since one of the syntypes is from the Seychelles Islands and the other from Mauri- tius; since Baker chose the specific epithet “sechellarum” he ob- viously had this Seychelles specimen in mind, and so it should be the lectotype. Therefore, I here designate Horne 194 (K), from Mahé, Seychelles, as lectotype, which means that this epithet replaces albobrunnea. The following new combination is thus necessary: Grammitis sechellarum (Baker) Morton (Polypodium sechellarum Baker in Hook. & Bak. Syn. Fil. ed. 2, 508. 1874). Those who wish to continue to recognize Xiphopteris as a distinct genus will have to make the appropriate new combination. Salvinia rotundifolia Willd. is recognized as adventive in the Lukungo River, Congo, but as I pointed out (Contr. U. S. Nat. RECENT FERN LITERATURE 125 Herb. 38: 75. 1967) this species has been misidentified by recent authors; the type in the Willdenow Herbarium is clearly a synonym of S. auriculata Aubl. I am not quite sure of the proper name for the species that has been called S. rotundifolia by Weatherby and others, but it may be S. minima Baker. Schelpe also records S. auriculata Aubl. from Lake Kariba, where it is a serious pest; however, I have had an intimation from workers who are studying this problem that the Lake Kariba plant is not really A. auriculata, but a sterile pentaploid, which would indicate that it is probably a recent hybrid that is spreading solely by vegetative means. Schelpe’s treatment of Elaphoglossum is going to be especially useful, because this is one of the most difficult genera throughout the tropics. The key is based almost wholly on scale characters, which is doubtless inevitable, since the species are so uniform in most ways other than the scales. Recent studies of the stipe anat- omy and spores indicate that there may be some additional usable characters. It is an open question whether Pteris quadriaurita Retz. is a species or an aggregate of many nearly allied species. Schelpe indicates that the question is not solved; in his treatment he is conservative and recognizes only one species, with four subspecies, which are not keyed out. Cheilanthes, one of the larger genera treated, is regarded in an inclusive sense, including Notholaena and Aleuritopteris, although Aspidotis is segregated, but Poly- podium is split up, following Ching’s lead, into Pleopeltis, Phyma- todes, Microgramma, Microsorium, and Polypodium. It should be mentioned that Schelpe recognizes Phymatodes K. B. Presl (Tent. Pterid. 195. 1836) as a valid genus, but in pub- lishing this generic name Presl cited as synonyms Dipteris Reinw. and Microsorium Link, both earlier validly published and legiti- mate genera. Therefore, by Art. 69 of the Code, Phymatodes Pres] was a superfluous name, and consequently illegitimate, since the earlier name Dipteris should have been adopted instead of a new generic name. Phymatodes is, by Art. 7, Note 4, typified by the type of the name that ought to have been adopted, namely by Dipteris conjugata; it is a straight synonym of Dipteris. The name 126 AMERICAN FERN JOURNAL Phymatodes could be used in Schelpe’s sense by conservation, but this hardly seems necessary. Its group is a small one, and is usually united with Microsorium, even by those who split up Polypodium, such as Copeland, who is by no means conservative in his recognition of genera in this group. Schelpe separates Phymatodes from Microsorium by the former having paraphyses in the sori and the latter not, but even if true this is not necessarily a generic character, and that it is always true remains to be dem- onstrated. Schelpe had only a single species of Phymatodes and only two, closely allied, of Microsorium. The numerous Malaysian species of these groups have hardly been eee Schelpe uses the spelling ‘‘M7zcrosorium,” as is usual and correct in my opinion. The name was originally spelled ‘‘Microsorum” by Link (Hort. Berol. 2: 110. 1833), and this spelling was adopted by Sledge in his treatment of the Polypodiaceae of Ceylon, but it was later changed by Link himself (Fil. Sp. 116, 135, 1841) to Microsorium. The Greek “‘soros,” generally Latinized as ‘‘sorus,” is masculine, and if Link had wanted to make a generic name from it he would have called it “Microsorus,” just as he proposed Cyclosorus and Camptosorus. Instead, he evidently intended a diminutive, the Greek “‘-ion,”’ Latinized usually as “ium,” on the analogy of Pteridium, and so the original ending ‘‘um”’ was properly corrected to “ium.”—C.V.M. A New Name For A Species or Potypopium From NoRTH- WESTERN Nora AMERICA, by Frank A. Lang. Madrofio 20: 53- 60. 1969.— ted the western Polypodium hespertum Maxon cytologically and Geode that there are two cytotypes, 4 diploid and a tetraploid, which can be distinguished morphologic- ally. The type of P. hesperium corresponds to the tetraploid; it has the sori oval and medial, the rhizome with a sweetish, licorice taste, and the rhizome scales without a dark median stripe. It oc- curs mostly in the interior, east of the Cascade Mountains. The diploid has the sori circular and submarginal, the rhizome acrid, and the rhizome scales often with a median dark stripe. It occurs mostly in the western mountains from British Columbia to the Sierra Nevada, but there are a few specimens from elsewhere Bane Recent Fern Lirera ture 127 (Wyoming, Colorado, Arizona). This diploid plant is called “Poly- podium montense F. A. Lang, nom. nov.,” and Polypodium amor- phum Suksd. Werdenda 1: 16. 1927, is cited as a synonym (Type: Suksdorf 11667). The holotype of P. montense is said to be Lang 211. This shows a confused concept, for if P. montense is a “nom. nov.” i.e. anew name for a species previously published then it must have the same type as the earlier name, that is, the type of P. montense would have to be Suksdorf 1166. Actually, since Lang gives a new Latin description and cites a new and different type, his species is apparently really considered a new species. However, it is in my opinion a superfluous name, since the legitimate species P. amor- phum Suksd. is cited as a synonym and that name should have been adopted. Lang’s reason for rejecting P. amorphum is that he considers it a monstrosity to be rejected under Art. 71 of the Code. He defines a monstrosity as ‘‘a plant that deviates greatly from the natural form of character, is abnormal, or is malformed,” but this is by no means what the Code means by a monstrosity. A true monstrosity would be something like Spondylantha Presl, which was based on a plant of Cissus sicyoides that was so trans- formed by a smut that it was not recognizable as a Cissus at all. But Polypodium amorphum is not like that at all: it merely has somewhat abnormal leaves but they are recognizable as belonging to Lang’s species since it is cited as an unquestionable synonym. Many species have been described from somewhat abnormal speci- mens, e.g., Polypodium trifurcatum L., which was based on a plant with the fronds forked at the apex; the name obviously applies to a common West Indian species. So far as I know no plant with the leaves forked at the tip has ever again been found, yet no one has ever questioned the validity and legitimacy of the name P. trifurcatum. Therefore, the name P. montense must be rejected as a superfluous new name for P. amorphum Suksdorf.—C.V.M. FLora PALAESTINA, PART ONE, TEXT, by Michael Zohary. Israel Academy of Sciences and Humanities, Jerusalem, 364 pp. + 2 maps. 1966.—This flora, which includes the pteridophytes of Israel, is not new but has only recently come to my attention. As might be imagined from its aridity, Israel is not a ‘“ferny 128 AMERICAN FERN JOURNAL country. In fact, there are only 15 species of pteridophytes that grow there. Any Fern Society members visiting Palestine will be able to identify the ferns they see merely by recognizing the genera, for each genus is represented usually by only one species. As a matter of fact, the families are so finely split that there is mostly only one species in each. The species are: Equisetum tel- mateia, E. ramosissimum, Ophioglossum lusitanicum, Cheilanthes ans, C. catanensis (Notholaena vellea), Adiantum capillus- veneris, Piss vittata, Anogramma leptophylla, Thelypteris palus- tris, Asplenium adiantum-nigrum subsp. onopteris, Ceterach offici- narum, Phyllitis sagittata, Dryopteris villarii subsp. australis, Polypodium vulgare var. serratum, and Marsilea minuta.—C.V.M American Fern Society New Members Mr. Frederick J. Ashby, 3219 Lorraine Ave., Kalamazoo Mich. 49001 Mr. Edward L. Bennett, Granby Florists, 345 Salmon Brook St., Granby, onn. Mrs. D. J. Burns, 1012 Irving Road, Birmingham, Ala. 35209 Mr. Hunsdon R. Cary, P. O. Box 415, Ojai, Calif. 93123 Mr. Dara E. Emery, 517 Junipero St., No. 2, Santa mipebec —— 93105 Dr. H. R. Fletcher, Royal Botanic Carden, Edinburgh, S and Mrs. David W. Miner, 1406 Alice Lane, Sulphur, La. oe Mr. H. L. Rachlin, 35 Park Avenue, New York, N. Y. 10016 Mr. Gregory L. Randolph, 4713 S. 100 E, Ogden, Utah 84403 Miss Sylvia Jane Rigby, Apt. 302, 105 Water St., Guelph, Ont., Canada Mrs. Charles B. Robinson, 2042 Pinehurst Street, Sarasota, Fla. 33581 Hh = G. rt gc 1804 E. 27th St., Jacksonville, Fla. 32206 , 2-2-5 Shimodo-Machim Kamo-Gun, Shizuoka-Ken, 415, Japan Mr. Sadao Sawai, No. 18, 1, Chome, Hiokacho, Chigusaku, oe Japan : Mire. d. Sickel-Zdzienick, 7927 Michigan Ave., Oakland, Cal 5 Mrs. Samuel R. Simpson, Sr., Hidden Gardens, Route 1, a en Franklin, N. C. 28734 Miss Lanora Lee Smith, Route 1, Box 941R, Porter, Texas 77365 Mr. Sarvjit L. Soni, 42 Delaware Ave., Toronto 173, Ont., Canada Mrs. Roger Tabb, 3055 8.W. Underwood Dr., Portland, Oregon 97225 Mrs. Peter Taylor, 917 Rugby Road, Charlottesville, Va 03 Mr. Raymond E. Pod Box 1980 Harrison Hall, Purdue ce. , W. Lafayette, Ind. 4 Mrs. Leslie D. ie 10501 N.E. 47th Place, Kirkland, Wash. 89033 Miss Irene Webster, P. O. Box 575, Hannandale, Fla. 33009 Exotic and Hardy Ferns Begonias BOLDUC’S GREENHILL NURSERY 2131 Vallejo Street St. Helena, California 94574 Open Saturdays and Sundays from 10 A.M. to 4 P.M. and by appointment Phone $63-2998—Area Code 707 Mail orders accepted CA oS VoL. 60 OctToBER-DEcEMBER, 1970 No. 4 American Fern Journal A QUARTERLY DEVOTED TO FERNS Published by the AMERICAN FERN SOCIETY ts DAVID B. LELLINGER C. V. MORTON ROLLA M, TRYON IRA L. WIGGINS CONTENTS A Major North American Range Extension for the — Spleenwort, Asplenium septentrionale..............---+--°-+: vip L. Emory 1294 Hard Water as a Limiting Factor in the Distribution of Isoétes echino- spora LeRoy Lee anp Frances Betknap 134 ee Indian Pteridophytes Used in Folk Remedies...........--. H.S.Purt 137° The Rhizome Scales of Platycerium......... Barpara Joe Hosuizaxr 144 Shorter Notes: Stem Cross-section Prints Aid in Identifying Horsetails; * Psilotum —— Spreading Northward;-Terminology of the — angial Structures of Equisetum; Kinds of Cystopteris in California. . 161 em Oa We a id cc Ra he EN wee CIs 6OS Sw Rae ots 165 Recent Form Literature... .c.c. 0 sees eevee e scene ccseeeeresescesee 167 American Fern Society. ........ 0... 0 ese c cee r eee ener en eeeenncees 169 es OO Wn io is vivn bce apres pee cere 8 oe ere eee nee eee 170 AGA 0s ae RING eee ee nd Cae e Sane ene ap cas inom (oat ee ‘ M } la i +t j al An? Che American Fern Society Council for 1970 ies Evans, 1g, Deparent of Botany, University of PE baa fms or K. hagrn Division of Plants, Carnegie Museum, Oe Penn- fagotto re Davmw B. Letuinaer, Smithsonian Institution, Washington, D. C. 20560. Editor~ in-Chief ational Society Representatives Warren H. Wacner, Jr., hee ar of — A.A.A.S. Representative Rotua M. Trron, JR., Harvard Uni A.I.B.S. Representative American #4 Fournal EDITORS Davin B. on eee Smithsonian Institution, Washington, D. S 20560. Ahi ¥ > OOO sic cs kee Smithsonian Institution, Washington, D. C. 20560. y Herbarium, Harvard University, Cambridge, Mass. 0213 Tra L. Viaea: ..-.Dudley Herbarium, Stanford University, Stanford, Calif — quarterly devoted to the — B10 fim Ave “a owned by the American Fern Society, and published at Baltimore, Md. 21211. § Second-clas postage paid a t Baltimore ‘of the Jour- nal are open to ne igee 2b rag hep heenangs 5 Fa ; membershi lists icharcnted taining specimens from different localities Matter for sobliaticn aud be addressed to the Edito: ges of ae, nid be addrewed t membership, eibeceiptiaan: orders Subscriptio: ns $5.50 gross $5.00 net (agency fee $0.50); sent free to m bers of the American can Fern Society (annual dues, 5,00; sustaining mem mberahi, Pea gees 2 $100.00). Extracted reprints, if ordered in advance, will be furnished a barn plus Back volumes $5.00 to $6.25 each, ingle back numbers of 64 singl —. $1.25 ciel oy ope each; over 80 pages, $2.50 each; Cumu oe dex to Volumes 50 psy Ten percent docsent on orders of six volumes Library and Bervarium Dr. pits H. Wagner, Jr., Department of Parrond University of Michigan, Arbor, Michigan 48104, is Librarian and Curator. Members —s borrow books and specimens at any time, the borrower paying all pin Mr. Neill D. Hall, oa an . 1225 Northeast 95th Street, — poe agit is Director. Spores exchanged and collection lists sen Gifts and Gifts and bequests to eee taekes © peed aries Se bers and to others interested in ferns. Herbarium — specimens, bo’ back issues ofthe Joona, and cash or other gifts are alae weloomed sre tax-deductible. Inquiries should be addressed to American Fern FJournal Vou. 60 OcTOBER—DECEMBER, 1970 No. 4 A Major North American Range Extension for the Forked Spleenwort, Asplenium septentrionale Davin L. Emory! The ‘“‘Forked Spleenwort,” A splenium septentrionale (L.) Hoffm., is a very unusual Spleenwort and one of the most striking ferns in the American flora. Presumably because of its peculiarly attenuated leaves (Fig. 1), the plant was recently referred to by Darling, (1964, p. 200) as an ‘“‘oddity.” Nevertheless it is a very wide ranging plant. Its distribution in western North America has been sum- marized by Lang (1969), who found the species recently for the first time in Oregon. Because of its ‘‘striking resemblance to tufts of grass” and its tendency to occur in very localized populations, it probably occurs in other as yet undiscovered places in the western United States; at present it is known there from the eastern slope of the Rockies from South Dakota and Wyoming south to western Oklahoma, New Mexico, Arizona, and Baja California. In the Old World, Broun (1938) reports it from the British Isles, Europe, northern Asia, and the Himalayas. Of the seven species of Spleenworts known in Britain, Manton (1950, p. 98) writes that ‘The rarest is Asplenium septentrionale . . ., a plant of south- ern affinities only found in a few presumably relict localities in the mountains of England, Scotland, and Wales.”’ The habitat given by Broun is ‘‘sheltered crevices in cliffs of igneous rocks.” During the second term of the 1970 summer session at Mountain Lake Biological Station of the University of Virginia, I was enrolled in the Pteridology course which has been taught there since 1961 by Dr. Warren H. Wagner, Jr., of the University of Michigan. A major emphasis in this course is upon student research in the field and laboratory. Accordingly, on August 2, 1970, a committee from 1 I wish to thank Mrs. Carolyn Crump for the photographs and Dr. Florence S. Wagner (working under NSF Project GB-8113) for the chromosome obser- vations. Volume 60, No. 3, of the JourNax, pp. 89-128, was issued Sept. 16, 1970. AMERICAN FERN JOURNAL VouumeE 60, Puate 16 Buurr in Monroz Co., West VIRGINIA, WHERE A. SEPTENTRIONALE GROWS RANGE EXTENSION FOR ASPLENIUM 131 the class, consisting of Carolyn Crump, Ronald and Shirley Fortney, Lewis Morgan, and me, began a pteridological survey of Monroe County, in southeastern West Virginia. This county, which is adjacent to Giles County, Virginia (the location of the Biological Station), seemed especially promising for field explora- tion because the class had already visited locations there discovered by E. T. Wherry during the late 1930’s for Phegopteris connectilis and Lycopodium porophilum, which are growing at this southern latitude at unusually low elevations for these northern plants. While we were climbing on the upper slopes of a bluff of loose, strongly brittle shale above a stream (Plate 16), we noticed what appeared to be several tufts of a small grass or sedge growing from crevices in the exposed, brightly sunlit, and extremely dry shale. In color, size, and texture, the narrow leaf tufts also seemed quite similar to bunches of pine needles. Closer examination revealed, however, that these ‘‘needles,” some of which were four inches long, were actually slightly expanded near their tips and forked. They bore on their undersides unmistakable fern sporangia. Hav- ing seen Asplenium septentrionale on the American Fern Society Colorado Foray in 1964, I quickly realized that we had made a remarkable discovery. Consultation of the literature indicated that our plants were nearly 1,200 miles from the nearest known station in western Oklahoma. When Dr. Wagner and the entire class visited the area on the following day, we counted sixty plants from four to nearly 40 feet above the stream level on the northwest-facing bluff. Diligent Searching up and down the stream revealed five more plants on another cliff about one-fifth of a mile upstream. One of the latter plants is shown in Figure 1, growing from the same crevice as A. platyneuron. As these plants on both cliffs were of all ages, the populations are definitely reproducing. Unless some calamitous disturbance takes place, the ferns should be able to maintain themselves indefinitely. We have no idea how old the populations are. It is entirely possible that other stations will be discovered, as there are numerous similar habitats nearby. AMERICAN FERN JOURNAL VotuME 60, PLaTE 17 4s A. PLATYNEURON (BELOW). Fig. 2. Curomosomes or A. SEPTENTRIONALE AT MEIOTIC METAPHASE. Fig. 3. SAME, CAMERA LUCIDA INTERPRETATION. RANGE EXTENSION FOR ASPLENIUM 133 Because of its great disjunction and the possibility that the local populations have become differentiated, we were especially inter- ested in determining the chromosome number. Manton (1950, p. 98ff.) reported that A. seplentrionale in Great Britain is tetraploid (n = 72 pairs at diakinesis). Cytological preparations of the plants we obtained, made by Dr. Florence 8. Wagner, showed unmistake- ably the same number (Figs. 2, 3). The “‘Septentrionale Bluff,’”’ at an elevation of about 2100 feet, is shown in Plate 16. It supports only a sparse vegetation. Trees include Pinus virginiana, P. strobus, Quercus prinus, Q. ilicrfolia, Betula lenta, and Robinia pseudoacacia. Among the shrubs are Vaccinium vacillans, Kalmia latifolia, Rhus radicans, and Parthe- nocissus quinquefolia. The herbs Aquilegia canadensis, Houstonia longifolia, and the attractive Campanula divaricata are in evidence here, the last in bloom at the time of our discovery of the fern. The loose shale is extensively covered by a variety of lichens (especially such fruticose types as Cladonia rangiferina) and a few mosses, including Polytrichum spp. and Leucobryum glaucum. Ferns associated with the Forked Spleenwort are Asplenvum trichomanes and A. platyneuron. In Great Britian and Europe A. septentrionale hybridizes with the first to yield A. X germanicum (Manton, 1950, p. 100ff.). This is the first time that A. septentrio- nale and A. platyneuron have been reported growing together. No hybrid plants of the latter parents have yet been discovered. A number of fronds collected from several plants on both days of our visits (Emory 70019, 70020) have been distributed to the following herbaria: US, GH, NY, MICH, WVA, and TENN. More precise information on the location in Monroe County, West Virginia, is not being published in the hope that the population can be protected. I do not feel that there is a need for oversampling, and I wish to urge the greatest caution in maintaining these plants in their natural state without disturbance. This report of Aspleniwm septentrionale in the Appalachian region constitutes a new addition to the flora of the eastern United States. The significance of disjunct eastern localities for western fern species will be discussed in another paper. 134 AMERICAN FERN JOURNAL LITERATURE CITED Broun, M. 1938. Index to North American ferns. 217 pp. Science Press, Lancaster, Pa. Daring, T. 1964. Southwest Vacation. Amer. Fern J. 54: Lana, F. A., 1969. The first record a Asplenium ae = ) Hoffm. in Or regon. Amer. Fern J. 59: 45-47. Manton, Irene. 1950. Problems of Gan Om and Evolution in the Pterido- phyta. xi + 316 pp. Cambridge Univ. Press. Mercerspure AcADEMY, MERCERSBURG, PENNSYLVANIA 17236 Hard Water as a Limiting Factor in the Distribution of Isoétes echinospora _ LeRoy Lree anp FraNcES BELKNAP Isoétes in Wisconsin has been shown by Swindale and Curtis (1957) to be a typical member of the soft water flora. Tryon et al. (1953) show it distributed in soft water areas, but absent from areas where hard water lakes are common. Moyle (1945) reports the Quillworts in Minnesota as members of a flora limited to a single chemical type: the soft water lakes found in the northeastern portion of the state. Although soft water is the typical location of Quillworts, they have been found in hard water. Lee reports Isoétes echinospora from Moshawquit lake. This Wisconsin lake has a specific con- ductance of 173 micromhos/em and a total alkalinity of 98 mgm/I CaCO:, which exceeds the upper limit of tolerance as reported by Moyle (1945). Seddon (1965) reports it in two hard water lakes in Wales, Llyn Llygerian and Llyn Llywenan. Music: (1945) suggests that the aquatic species of Isoétes are _ found in soft water because they can not tolerate a total alkalinity DISTRIBUTION OF [SOETES 135 of above 50 mgm/l CaCOs;, but since they have been reported from other than the usual soft water sites, the distribution can not be completely attributed to water chemistry. Seddon (1965) con- cluded from field observations that competition is responsible for the restriction of I. echinospora to soft water. o determine if Quillworts could survive and grow in hard water without competition, I. echinospora var. braunii (Dur.) ngelm. was grown under laboratory conditions of varying water chemistry during one summer growth period. TaBLe I. Water ANALYSIS OF EXPERIMENTAL Tanks! total alkalinity . conductance water color (CaCO; mgm/l) pH (micromhos/cm) (PI-Co units) Crooked Lake 29 15 119 15 Crystal Lake 92 7.6 231 da Wis. River 112 S.2 282 65 Fish Lake 148 7.9 304 35 Pretty Lake 160 8.3 483 40 Lake Ripley 184 8.6 431 20 i 228 738 643 38 8.0 585 30 Merrill Springs 281 1 Water analysis furnished by Mr. Ronald Poff, Water Classification — ist, Wisconsin Department of Natural Resources, Madison, Wisconsin Eight 10-gallon aquaria were filled with water and substrate, each from a specific body of water. Water sources were chosen to provide a range from very soft to very hard water (see Table I). Experimen- tal plants were collected from Crooked Lake, Burnett County. For each tank, ten plants were randomly selected from the collee- tion, weighed, the number of leaves counted and measured, and then planted. Twelve hours of light was supplied daily by Gro-lux fluorescent bulbs to provide 100 ft-c of illumination at the water surface. Weekly water analyses determined that little change in water chemistry occured over the experimental period. 136 AMERICAN FERN JOURNAL At the end of 21 weeks all plants were still surviving in the soft water control (Crooked Lake) and the hard water aquarium from Ripley. Seventy per cent of all the experimental plants survived; they grew an average of 5 new leaves and had 2 cm of leaf and root growth, the same as the average in the soft water control tank. Sporulation did not occur. At the termination of the experiment the plants were replanted, additional hard tap water (280 mgm/] CaCO;) was added to each tank to replace evaporation loss, and continuous illumination at a high intensity was provided. After five months all plants still survived and appeared healthy. Several were sporulating. ile the results of this experiment are tenuous due to the death of 30% of the experimental plants, they do support the belief that water hardness is not the single major limiting factor in distribu- tion. The continued growth and sporulation with increased illumi- nation, plus the fact that the two tanks which had 100% survival were significantly less colored than the others (see Table J), indicates that the amount of light reaching the plants may be an important factor in the distribution of Isoétes echinospora var. braunit. LITERATURE CITED Moxy:g, J. 1945. Some chemical factors influencing the distribution of aquatic plants in Minnesota. Amer. Mid]. Nat. 34: 402-420. Seppon, B. 1965. Occurrence of Isoétes echinospora in eutrophic lakes in Wales. Ecology 46: 747-748. Swinpate, D. and J. Curtis. 1957. Phytosociology of the larger submerged plants in Wisconsin lakes. Ecology 38; 397-407. Tryon, R., N. Fassert, D. Duntap, and M. Diemer. 1953. The ferns and ferxi allies of Wisconsin. 158 pp. University of Wis. Press, Madison, Wis. Mapison Pusiic ScHoots, Mapison, Wisconsin 53715. PTERIDOPHYTE FoLtK REMEDIES 137 Indian Pteridophytes Used in Folk Remedies H. 8. Purr The medicinal qualities of ferns, real or imaginary, are mentioned as early as 300 B.C. by the Greek philosopher Theophrastus (Corne, 1924a) and by his Indian contemporaries Sushrut and Charak. Dioscorides also speaks of brake fern, male fern, and others. Among the medicinal plants mentioned in old Indian texts, two are def- initely ferns. One of these, called ‘‘Myurshikha” or ‘“‘Myurpankha,” meaning sees feather, has been identified as Adiantum cau- datum L. by some persons; others consider it to be sag radiata Swarts Link. The other drug called “Hanspad” o “‘Hansraj,”’ meaning swan’s foot, appears to be some species of Adiantum, most probably A. capillus-veneris L., A. lunulatum urm. f. or A. venustum D. Don. In addition to these plants there are about 40 species of pteridophytes used as drugs. Of these Dryopteris filiz-mas (L.) Schott is included in the pharmacopoeias of many countries. In the 1955 Pharmacopoeia of India this plant is replaced by the allied species, D. odontoloma (Moore) C. Chr., D. marginata (Wall.) Christ, and D. schimperiana (Hochst.) C. Chr., and in the 1966 Pharmacopoeia of India by D. odontoloma, D. marginata, D. chrysocoma (Christ) C. Chr., D. ramosa (Hope) C. Chr., and D. barbigera (Hook.) Kuntze. Lycopodium clavatum is caclided * in the Indian Pharmaceutical Codex (Mukerji, 1953) and in the proposed Indian Homeopathic Pharmacopoeia. Equisetum arvense L. is included in the U.S.S.R. Pharmacopoeia (1961) and in the German Pharmacopoeia (Wealth of India), and Puri (1969) has recommended its inclusion in the Indian Pharmacopoeia. The species of Adiantum mentioned above are used unofficially in some of the patented cough preparations marketed in India. The rest of the plants mentioned below are used in domestic and rural medicines. 1 The author is peety indebted to Prof. P. N. Mehra for his keen interest and to Mr. C. V. Morton for his kind suggestions during the preparation of this article. 138 AMERICAN FERN JOURNAL Actiniopteris radiata (Swartz) Link—Used as an anthelmintic (Watt, 1889; Kirtikar & Basu, 1935; Chopra et al, 1958; Chunekar & Pandey, 1969), as an alterative in prolonged malarial fevers (Nadkarni, 1954; Chopra et al.), and as an astringent to arrest hemorrhages (Chunekar & Pandey). Adiantum aethiopicum L.—Infusion of leaves used as an emol- lient in coughs and diseases of the chest (Caius, 1935). Adiantum capillus-veneris L.—According to Watt, the bulk of Adiantum sold medicinally in India is this species. A decoction of the fronds of this plant is considered quite effective in all types of bronchial troubles (Watt; Nadkarni; Steinmetz, 1954; Wren, 1956; Uphof, 1959). An infusion of the herb serves as a shampoo against dandruff and also promotes hair growth (Watt; Steinmetz; Wren). The plant is considered as an emmenagogue (Nadkarni; Uphof). The juice of the plant with pepper is recommended for all types of fevers (Nadkarni; Biswas, 1955). It is used for hydro- phobia by the physicians of the Persian system of medicine (Watt). Adiantum caudatum L—Fronds used for diabetes, coughs (Kirtikar & Basu; Chopra et al), and for migraine (Stewart, 1869). Also used externally for skin diseases (Chopra et al). Adiantum lunulatum Burm. f—One of the constituents of Hansraj, the drug esteemed in India for coughs. It is considered a bronchio-dilator, diuretic (Chopra et al) and pectoral (Caius). In western India used extensively in the treatment of fevers of children; the rootstock is considered good for fever and elephan- tiasis (Kirtikar & Basu). Adiantum venustum D. Don—According to Kirtikar and Basu the fronds of this plant are the source of the Indian drug Hansraj. Because of the black color of the stalks of the fronds, it is some- times known by the name of black Hansraj. The plant is adminis- tered as an anodyne in bronchitis and is considered a diuretic and emmenagogue (Stewart, Kirtikar & Basu; Chopra et al). It is a tonic, febrifuge, and expectorant, and is particularly useful as a tonic during convalescence from fevers (Caius). Asplenium adiantum-nigrum L.—A decoction or syrup of the fronds is used as an emmenagogue (Kirtikar & Basu). It is bitter, PTERIDOPHYTE FoLtK REMEDIES 139 diuretic, and laxative, and is considered useful in ophthalmia and diseases of the spleen and jaundice. It is believed to cause sterility in women (Chopra et al.). Asplenium falcatum Lam.—Used in enlargement of the spleen, incontinence of urine, calculus, jaundice, and malaria (Kirtikar & Basu Asplenium ruta-muraria L.—It is used for rickets (Chopra et al), and as an expectorant (Nadkarni) and deobstruent (Kirtikar & Basu). According to Steinmetz, ‘“Herba Rutae Murariae” is an expectorant and cures diseases of the spleen. It prevents hair from falling and makes it grow (Fernie, 1914; Steinmetz). Asplenium trichomanes L.—Considered a laxative (Fernie; Chopra et al) and anthelmintic (Nadkarni). The leaves are sweet, mucilaginous, and are an expectorant highly useful in pulmonary disorders (Caius). Athyrium filiz-femina (L.) Roth.—Used as a substitute for the Male Fern, but has been found quite ineffective (Chopra et al). Botrychium spp——Various species of this genus, such as B. lunaria (L.) Swartz, B. ternatum (Thunb.) Swartz (Chopra et al), and B. virginianum (L.) Swartz (Kirtikar & Basu), are used in dysentery, ruptures, and for healing wounds (Wealth of India). Cibotium barometz (L.) J. Smith—The rhizome is used as a vermifuge and the root as a tonic. The long, silky, yellowish hairs at the base of the fronds cause rapid coagulation of the blood and when properly used are useful for arresting hemorrhages from capillaries (Wealth of India). The root is employed as a tonic in China, where it is said to exercise a special action on the genito- urinary organs (Kirtikar & Basu). Used in typhoid and in the treatment of dyspepsia and coughs (Chopra et al). Drynaria quercifolia (L.) J. Smith—The rhizome is bitter, tonic, and astringent to the bowels. It is also used in typhoid and in the treatment of dyspepsia and coughs (Chopra et al). A decoc- tion of the rhizome is used as an astringent (Uphof). Dryopteris filiz-mas (L.) Schott——One of the oldest anthelmin- tie drugs, commonly known as Male Fern, and used since ancient . 140 AMERICAN FERN JOURNAL times for expelling tape-worms from the intestines. This fern does not grow naturally in India, but various allied species which are up to British Pharmacopoeial and United States Pharmacopoeial standards are used (Indian Pharmacopoeia, 1955, 1966). The use of D. filiz-mas does not appear to be harmless. According to Steinmetz, its application may cause paralysis of the muscles and nerves, and even blindness. This may be due to the presence of the enzyme thiaminase, which destroys thiamin (Pohl, 1955). Equisetum arvense L.—‘‘Herba Equisiti Arvensis” has diuretic and astringent qualities (Steinmetz; Wren) and is quite useful for urinary and bladder diseases (Uphof), and for stopping the flow of blood in urine (Steinmetz). An oral administration produces a decided increase in blood corpuscles. The plant is used for acidity of the stomach and dyspepsia. (Wealth of India). Equisetum debile Roxb.—Diuretic and astringent (Biswas), and given for gonorrhoea (Stewart). Helminthostachys zeylanica (L.) Hook—The plant has intoxi- cating and anodyne properties and is used in sciatica (Chopra et al; Wealth of India). Rhizome used for malaria (Uphof). The Malays regard the rhizome as a tonic, and eat it with betel for whooping cough. In Java used for dysentery, catarrh, and the early stages of phthisis (Wealth of India). Lycopodium cernuum L.—In Malaya a decoction of the plant used as a lotion in beri-beri and also for coughs and uneasiness in the chest. An embrocation of the ashes in vinegar is recommended for skin eruptions (Wealth of India). Lycopodium clavatum L.—Mainly the spores of this plant are used in medicine. English druggists included this powder in their list of drugs before 1692. It was introduced in the U.S. Pharma- copoeia of 1860 (Whitebread, 1941). This drug has been used internally with good results in urinary disorders, such as the spasmodic retention of urine, catarrhal cystitis, and chronic kidney diseases causing pain in the kidney, ureter, and bladder (Wren; Nadkarni; Biswas). Also used in dyspepsia, catarrhal gastritis, and as a general and gastric sedative (Wren). The plant is also given to stop hemorrhages after childbirth (Biswas). In the form PTERIDOPHYTE FotK REMEDIES 141 of a tincture it is given for rheumatism, epilepsy, and pulmonary disorders (Nadkarni). An aqueous extract of it has been employed in various countries as an antipyretic (Wealth of India, 1962). Sometimes the spores of L. annotinum L., L. complanatum L. and L. selago L. are also used for the above purposes (Wealth of India). Frequently Lycopodium spores are adulterated with such substances as slightly roasted and colored starch, dextrin, and various pollens (Whitebread). The Lycopodium powder has side effects. Whitebread has quoted instances where it has acted as foreign body and has given rise to an indolent chronic post-operative inflammatory reaction. The alkaloids of this drug have also been found harmful to warm- blooded animals. A variety of reactions, such as pressor effects, stimulation and contraction of the uterus, and paralysis are at- tributed to these alkaloids (Wealth of India). Lygodium circinnatum (Burm. f.) Swartz—The spikes of this plant are reported to be used in Indonesia as an external applica- tion for wounds (Wealth of India). Lygodium fleruosum (L.) Swartz—Used as an expectorant. Fresh rootstocks are applied externally for rheumatism, sprains, scabies, eczema, and cut wounds, and are reported to be particularly useful for carbuncles (Caius). Lygodium japonicum (Thunb.) Swartz—Used as an expectorant. In China a decoction of the vegetative parts and spores is used as _ a diuretic (Wealth of India). Lygodium microphyllum (Cav.) R. Brown—A decoction of leaves given for dysentery. Leaves are also applied as poultices for skin diseases and swellings (Kirtikar & Basu). Marsilea quadrifolia L—This is probably the plant used in ancient India under the name of “Sparka” for leprosy and skin diseases, fever, and for poisoning of blood (Chunekar & Pandey). Ophioglossum vulgatum L.—The leaves boiled in oil or fat are considered a remedy for wounds. This preparation cools inflam- mation (Fernie; Chopra et al). Osmunda regalis L.—Considered a tonic and styptic, and used against rickets (Chopra et al). The middle part of the plant boiled 142 AMERICAN FERN JOURNAL in liquid is considered good for wounded persons. The rootstock, stamped in water or gin until the liquor becomes a stiff mucilage, has been used to cure back pains (Fernie). Polypodium vulgare L.—An infusion of ‘“Herba Polypodii” is a mild laxative and expectorant. An aqueous extract of ‘‘Rhizoma Polypodii” is recommended in the obstruction of viscera, liver and gland diseases, and coughs and chills (Steinmetz). In northern India this or a related species is used as an alterative (Stewart). Polystichum squarrosum (D. Don) Fée—The rhizome of this plant is sold in the Indian market under the name ‘‘Nirbissi,”’ and the drug is used as an antidote to poisons; it is a possible substitute for the male fern (Mittal, 1953). Pteridium aquilinum (L.) Kuhn.—The rhizome is astringent and is useful for diarrhea and inflammation of the gastric and mucous membranes. Boiled in oil or hog’s fat, the rhizome is made into an ointment for wounds. A decoction of the rhizome and fronds has been given in chronic disorders arising from obstruction of the viscera and spleen (Wealth of India). It is also used instead of hops (Corne, 1924b). Pteris ensiformis Burm. f.—The juice of the young plant is stated to possess astringent properties; a decoction of fresh fronds is given in dysentery. The juice of the rhizome is applied in the glandular swelling of the neck (Wealth of India). Pteris multifida Poir. (P. serrulata L. f.)-In China a tincture or decoction of rhizome is given in dysentery. Also said to be a good vermifuge. The toasted fronds and rhizome are made into a paste with sesame oil and applied to the skin afflictions of infants (Wealth of India). : Selaginella spp.—Apparently dry plants of some species appear to become fresh and green again when put into water. Because of this, physicians of the Indian system of medicine use it as a re- juvenator for the human body. It is also given in mental diseases, rickets, blood vomiting, and spermatorrhoea (Chunekar & Pandey). Tectaria polymorpha (Wall.) Copel—Used as an anthelmintic (Kirtikar & Basu). PreriDoPpHyTE FotK REMEDIES 148 LITERATURE CITED Biswas, K. 1955. Common Medicinal Plants of Darjeeling and Sikkim. otanic Garden, Calcutta. Caius, J. F. 1935. The Medicinal and Poisonous Ferns of India. J. Bombay Nat. Hist. Soc. 38: 341-361. Cuopra, R. N., Cuopra, I. C., Hanna, K. L., and Kapur, L. D. ee Tndigenoits Drugs of India. U. N. Dhur and Sons, Calcu CruunEKaR, K. C. and Panpery, G. S. 1969. Bhavparkash re ea (in Hindi). The Chowkhamba Viden Bhawan, Varanasi, India. Corng, F. E. 1924a. Ferns—Facts and fancies about them—II. Amer. Fern oan, Ibid. —III. Amer. Fern J. 14: 115-118. FEeRNIE, Ww. 7. 1914. Herbal eis ed. 3. John Wright & Sons, Bristol. Kirtrkar, K. R. and Basu, B. D. 1935. Indian Medicinal Plants, Part IV, ed. 2. Lalit Mohan Basu, Allahabad, India. Mirrau, T. C. 1953. Pharmacognosy and phytochemistry of some Indian ee of Polystichum and Athyrium. M. Pharm. Thesis, Panjab Univ., Chandigarh, India. Moxerii, B. 1953. Indian Pharmaceutical Codex, vol I. Council of Scientific and Industrial Research, New Delhi Napxarnl, K. M. 1954. Indian Materia Medica, vol I. Popular Book Depot, ombay. Pout, R. W. 1955. Toxicity of Ferns and Equisetum. Amer. Fern J. 45: 95-97. PHARMACOPOEIA OF INDIA. 1955. Manager of Publications, New Delhi. ————. 1966. Second edition. Manager of Publications, New Delhi. Puri, H. bs 1969. Scope of including some indigenous medicinal plants in I. P. Indian J. Hosp. Pharm. 6: 102-106. STATE ey RET, OF THE Unton oF Soviet Socraist Repusuics. 1961. 9th edition, Moscow seme E. F. 1 1954. Materia ora tne 9 E. F. Steinmetz, Kaizer- acht, Amsterdam, Nether ee ar J.L. 1869. Panjab seer ae t. Printing Press, Lahor Upuor, J. C. Tu. 1959. Dictionary of Economic Plants. Hafner, ie York. Warr, G. 1889-1892. Dictionary of Economie Products of India, Parts I-IV. . H. Allen. WEALTH a Inp1A. 1948-1969. Parts I-VIII. Council of Scientific and Indus- rial pemeerten New Delhi. aellees C. 194 1. Bewa ware of “Lycopodium.” Amer. Fern J. 31: 100-102. Wren, R.C. 1956. Potter’s New Cyclopaedia of Botanical Drug and Prepara- tion. Sir Issac Pitman and Sons, London Botany DEPARTMENT, PANJAB U ,C INDIA 144 AMERICAN FERN JOURNAL The Rhizome Scales of Platycerium! BARBARA JOE HosHIzAKI Considerable diversity exists in the structure of fern rhizome scales. Although the more conspicuous features of scales have long been used in describing and identifying some groups of ferns, the more detailed aspects have generally been ignored. Modern mono- graphs and treatments on ferns have contributed to a better understanding of the value scales may have in the interpretation of evolutionary relationships. The object of this paper is to de- scribe some of the rhizome scale features of Platyceriwm that seem to be helpful in determining the evolutionary trends within the genus. Some similar features have been pointed out by Wagner (1952, p. 77) in the genus Diellia. Before the rhizome scales could be studied for structures that might be of phylogenetic value, it was necessary to determine first the probable evolution of the Platycerium species. For this, I used a variety of characters other than those of the rhizome scales. Reliable characters included stele types, sporangial arrange- ment, nature of the sporangial stalk, and completeness of the annulus. Only after I had arrived at the probable evolution of the species did I examine the rhizome scales in detail to determine how their features correlated with the patterns I had determined. The three main lines of evolution are summarized in Table I. The Afro-American line includes P. andinum and two branches both starting from P. quadridichotomum. One branch ends in P. angolense and the other branch ends in P. ellisii. The sequence of Species in the other two lines is essentially linear, as listed. In selecting rhizome scales for study several factors must be kept in mind. The scales on individual plants vary, but species differences are usually consistent, providing the factors of the age of the plant, weathering, and selection of a typical scale from the sampling are taken into consideration. Immature or small plants _ _* Research supported by a National Science Foundation Science Faculty Fellowship. ; Pa ity RHIZOME SCALES OF PLATYCERIUM 145 generally have smaller, lighter, and fewer bicolorous scales bearing fewer hairs than those on mature plants. On fully mature plants scales may have suffered damage by weathering and must be assessed accordingly. The best scales for study are those that are starting to darken on full grown plants. Even in perfect material there will be variations; the largest, darkest scales usually show the characters of the species more clearly than the narrower, smaller, lighter colored scales or fibrils. In regards to selection of trichomes for study, Carlquist (1961, p. 33) aptly states, ‘“Each trichome type in each species shows some variability and thus each type should be studied as a population having extremes and a typical condition.” The plants used in this study are from cultivation; for most the specific place of collection is not known.? Silhouettes of the scales were made by tracing photographs; drawings of the hairs were made with the aid of a microprojector. Morpno.oey AND EvoLUTIONARY SIGNIFICANCE OF SCALES AND THEIR Hairs Scale distribution——Rhizome scales are most conspicuous on the buds, but also cover the entire surface of the stout rhizome and extend up the phyllopodia occasionally to just beyond the abscis- sion layer. On the bud the many light colored scales are tightly overlapped and appressed to the surfaces. Progressing to the older parts of the rhizome, the scales become darker, less appressed, and may twist or curl to form a thick, chaffy covering; they are per- sistent. General shape ——Each plant bears scales of various widths and lengths; the larger, darker scales are here considered the represen- tative scales of a species. For most species these are more or less narrowly triangular. However, the scales of P. andinum and P. angolense (Figs. 14, 15) are noticeably linear-triangular, and in P. 2 Messrs. Ed Franks, Rudy Ziesenhenne, Marcel Lecoufle, and John Roach very generously provided most of the plants and specimens for this study, and I am greatly indebted to them for their many kindnesses. I wish to thank ss also to wled. ) given : me by Dr. Mildred E. Mathias. The major part of this work was done at the University of California, Los Angeles, where facilities were kindly provided. 146 AMERICAN FERN JOURNAL coronarium and P. ridleyi they are obovate (Figs. 29, 30). All but P. coronarium and P. ridleyi have fibrils (very narrow scales) which also vary in width and length on the same plant; the nar- rowest are uniseriate flattened hairs and the widest are indistin- Tasie I. Species anpD THEIR RHIZOME SCALE CHARACTERS Apex Scales filiform narr (0) ) (0), Scales acule Character states- Fibrils inter- ; 3 = very specialized present mediate papery blunt 2 = specialized (0), (1), 0), (2), 1 = intermediate absent broad papery round 0 = generalized (2) (2) (2) (3) AFRO—AMERICAN LINE P. andinum 0 0 0 0 P. quadridichotomum 0 0 0 0 P. stemaria 0 0 0 0 P. angolense 0 0 0 0 P. madagascariense 0 0 0 0 P. vassei 0 0 0 0 P. elli 0 0. 0 0 JAVAN-AUSTRALIAN LINE P. willinckii 0 0 0 0 P. veitchii 0 0 0 0 P. bifurcatum 0 0 0 1 P. hillii 0 0 0 1 Mauayan-—Astatic LINE P. wallichii 0 0 0 0 P. holttumii 0 1 0 a P. wandae 0 1 0 2 QP, e 0 Z 2 3 P. coronarium 2 Pe 2 3 P. ridleyi 2 2 2 3 NNR Re oocoococo o guishable from narrow scales. The almost universal presence of fibrils in the genus seems to indicate the ancestral type had fibrils which developed into progressively broader scales. It is generally accepted (Bower, 1923, pp. 201-205) that hairs preceded fibrils and that fibrils developed into scales. Following this line of reasoning, RuHIZOME SCALES OF PLATYCERIUM 147 narrow scales probably gave rise to broad scales. Extremely broad scales (Figs. 26-30) seem to be linked to the development oF: flabelloid margins. The scale length varies from 2 to 35 mm, the longest being those of P. wandae and the shortest those of P. Hairs or Special- Basie Special- papillae ized hairs tized Flabelloid marginal hairs non hairs Partial margin (0), sub- none (0), (0), none or stellate none (0), marginal — semi- 1-7- mostly hairs narrow Total (1), (1), celled 2-5- absent (1), inter- value of super- _— mostly (1), celled (0), mediate special- ficial (2), all S-celled (0),1-2 present (2), ized (2) spec. (3) (2) (1), 1 (2) (2) broad (3) states 0 2 0 0 2 0 4- 0 1 0 0 0 0 1 0 1 0 0 0 0 1 0 2 0 0 2 0 4 0 1 0 0 0 0 1 1 2 0 1 0 0 4 2 3 0 2 0 0 ‘i 0 1 0 0 0 0 2 0 1 0 0 0 0 2 0 1 0 0 0 0 4 0 1 0 0 0 0 4 0 0 0 0 0 0 0 1 0 2 0 0 1 7 1 1 1 0 0 2 8 1 1 0 0 0 2 il 0 3 0 1 0 3 16 1 5 0 2 0 3 18 willinckii. The scales on the phyllopodia are larger than those on the rhizome. The texture of the scales is usually firm, but some FP grande scales are thin and papery and those of P. coronarium and P. ridleyi are also except at the center; this is probably a special- ized condition. 148 AMERICAN FERN JOURNAL Scale apex.—Scale apices may be long attenuate to acute, to blunt, rounded or emarginate (Figs. 2-5). Typically the attenuate apices end in a long, often gland-tipped filament. Acute apices have fewer and stouter cells in the filament; blunt apices are multicelluar and sometimes apiculate or emarginate. The apices may often be branched into a number of hairs or filaments. If it is true that scales have been derived from hairs, then the general- ized scale apex would be attenuate and filiform and the specialized apex blunt and multicellular. There would be a trend toward increasingly broad apices. There remains, however, the possibility that the extremely long attenuate apices of P. angolense and P. andinum may be a specialized condition. In the clear evolutionary line of the Malayan-Asiatic group there is a series from generalized to specialized apices. The inter- mediate state consists mostly of acute apices that are rarely fili- form. The Javan—Australian group has the generalized to inter- mediate state; the Afro-American group tends to the generalized. Scale base-—The scale base may be slightly narrowed, rounded, truncate, or subcordate to cordate (Figs. 9-13); however, most are unmodified. Most scales are sessile and attached by the length of their basal edge (Fig. 9), which may be a broadened area if the scale is bent and appressed to the rhizome (Fig. 10). Scales may also be subpeltate (pseudopeltate) to peltate. In all cases the scales interrupt the rhizome epidermis at their point of attach- ment. The subpeltate condition (Fig. 12), found on some scales of P. willinckii, P. veitchii, and P. bifurcatum, is often accompanied by a subcordate or cordate base; the attaching tissue is a long or _ Short flange. Platycerium hillii and some forms of P. bifurcatum have weakly peltate scales with rounded bases, which are most frequently found on the phyllopodia and which tend to detach easily (Fig. 13). There is a definite stalk or short flange which _ attaches to the rhizome in a slight epidermal depression. Peltate scales are generally considered more advanced than subpeltate or other unmodified ones (Bower 1923, p. 200). Scale ule margin.—The scale margin may be entire, or bear papillae or hairs, or be flabelloid (see the separate discussions on hairs an RHIZOME SCALES OF PLATYCERIUM 149 flabelloid margin). Small, dark protuberances are sometimes seen on or near the walls of the marginal cells of P. grande, P. wandae, and P. bifurcatum. They seem to be persistent cell nuclei which have not deteriorated, for in the younger parts of the scale nuclear stains bring out obvious nuclei which if examined in progressively older cells move toward the wall and appear as the small dark protuberances. Bicolorous scales —The central middle to basal areas of the scale are marked by a stripe on at least some scales of most species. The stripe is composed of cells which have all thickly pigmented brown to black walls. A scattering of lighter colored cells with only their lateral walls thickened may be found among the darker cells. As there is considerable inconsistency in color and pattern of these stripes, it is diffcult to use these to evaluate phylogenetic trends. The stripe extends to or nearly to the apex in members of the P. bifurcatum group more frequently than in other species. Platycerium grande often lacks a well defined stripe, particularly on the larger scales of the phyllopodia. Scale development.—Rhizome scales begin their development from uniseriate hairs. These hairs divide longitudinally, trans- versely, and clinally at the thickened center to form the scale. For cell details see Figure 1. In non-peltate scales the enlargement of the base is by transverse and longitudinal cell division. Peltate scale development in P. hillii often shows a cupping in or turning under of the basal auricles, which are eventually obliterated by thickening, enlargement, and overtopping growth of the whole basal area, resulting in a thick, rounded base (Fig. 13). The attaching cells that were between the auricles become submarginal and differentiate into a short flange or well-defined stalk. This scale development is similar to that illustrated for Pyrrosia (Nayar, 1961, figs. 7-15, p. 166), except that there are no auricles. Hair distribution —Hairs are found on all scales except those of P. wallichii. All others have hairs along the edge of the scales; the hairs are sparse near the basal edge. Platycerium bifurcatum and P. hillii tend to have long, tangled hairs near the apical margin of the scales, but this is not always consistent (Fig. 4). VoLuME 60, PLATE 18 AMERICAN FERN JOURNAL Fig. 1. P. ANGOLENSE, YOUNG SCALE, ig. 2. P. CORONARIUM, MATURE SCALE, M APEX. G: F- SCALES. GENERALIZED EXCEPT FOR HAIRS. SPECIALIZED, MARGIN PAPERY, FLABELLOID. Fig. 3. P. VEITCH, FILIFOR 5. P. GRANDE, BLUNT APEX. F PLATYCERIUM RHIZOME 1G. 5: 2, Fig. 4. P. HILLII, ACUTE APEX. HOLTTUMII, MARGIN WEAKLY FLABELLOID. Fig. 7. P. WANDAE, MARGIN FLA BEL- Low. Fig. 8. P. GRANDE, MARGIN STRONGLY FLABELLOID. RuHIZOME SCALES OF PLATYCERIUM 151 Hairs on the broad surfaces of the scales are found near the apex occasionally in P. angolense, and over much of the surface in P. ellisit and P. vassei (but more sparsely in the latter), and on the face of the marginal cells in P. ridleyi. The dense marginal fringe of hairs in P. holttumii, P. wandae, and P. grande scales (Figs. 6-8) arises from marginal and submarginal cells of the flabelloid margin. Having the hairs consistently scattered over the surface of the scale or spreading to a submarginal position occurs in most : advanced to moderately advanced species and is to be regarded — aS more specialized than having hairs limited to marginal cell | edges. In the apparently related genus Pyrrosia, a similar distribu- tion of hairs is also found among the advanced or moderately advanced species (Nayar, 1965, p. 13) Hair types—The hairs found on the rhizome scales fall into three intergrading types: glandular, ray, and basic. The glandular types are typically stout, round at the apex, and contain a dark, gland-like substance. The ray types appear much like the rays of the stellate hairs of the lamina; most are whitish and blunt to pointed at their apex, and generally flat, although some are conical ~ and often a bit darker. Basic hairs are close to the ray type in : appearance and color, but are typically bent, twisted, or crooked, and not as flat and white as the ray type. Shorter basic hairs bear a close resemblance to the marginal papillae, whereas longer ones may be setiform. Typical glandular hairs are found in P. ridleyz (Fig. 30), typical ray types in P. vassei (Fig. 198), and basic types in P. grande (Fig. 288). Some basic types may blend into the ray type, a common condition in P. wandae (Fig. 27). All three kinds of hairs and their intermediates are found in the Malayan—Asiatic line (Figs. 25-30). The Javan—Australian line (Figs. 21-24) and the more primitive members of the Afro-American line (Figs. 16-18) have mostly basic to glandular types. Basic to glandular inter- mediate hairs tend to have the contents of the cell collapsed, causing a distortion of the cells (Fig. 17). Origin of the hair types—The presence of hair-like papillae and — their similarity to basic hairs suggest that basic hairs are derived from elongate papillae. This follows the dictum that trichomes AMERICAN FERN JOURNAL 20 SWS bia/ V7 al eer \ d 4k W Puatycerium Raizome ScaLes anp Hairs. Figs. 9-13. RHIZOME SCALE BASES, NOT TO SCALE. Fic. 9. P. sTeMARIA, BROAD ATTACHMENT. Fia. 10. P. VASSEI, BROAD ATTACHMENT, SCALE BENT. Fig. 11. P. wa ANDAE, SUBCORDATE BASE. Fic. 12. P. WILLINCKIT, SUBPELTATE BASE. Fig. 13. P. HILLII, PELTATE BASE. Figs. 14-30. RuizoME SCALES AND HAIRS SHOWING CHANGES ALONG EVOLUTIONARY LINES. Figs. 14-20. ie LINE. Fig. 14. P. ANDINUM. Fig. 15. P. ancouense. Fig. 16. P. sremarta. Fic. 17. P. QUADRIDICHOTOMUM. Vouume 60, Pate 19 Fig. 18. P. MaDAGASCARIENSE. Fig. 19. P. vassEI. A, MozaMBIQuE; B, Mapa- GASCAR,; C, CULTIVATED. Fig. 20. P. Evuisit. Fias. 21-24. JavAN-AUSTRALIAN LINE. Fig. 21. P. winzincxn. Fig. 22. P. verrcuu. Fig. 23. P. srruRCATUM. Fig. 24. P. nmin. Figs. 25-30..Matayan-Astatic Line. Fig. 25. P. wat LICH. Fig. 26. P. nourrumu. Fig. 27. P. wanpaer. Fic. 28. P. GRANDE HAIRS FROM MEDIAL MARGIN; B, HAIRS FROM BASAL MARGIN. Fig. 29. P. coro- NARIUM. Fig. 30. P. rmieyi. The abbreviation is: M = MARGINAL CELLS. 154 AMERICAN FERN JOURNAL originate from papillate epidermal cells (Netolitzky cited in Carl- ar 1961, p. 34). This hypothesis is supported by the presence n P. holttumii and P. grande of very short to long papillae inter- aed with basic hairs, which are distinct only by the cell wall separating the hair cell from the trichoblast. Uniseriate hairs are generally regarded as the basic type from which many other variants are derived (Wagner, 1964, p. 90). The origin of the glandular hair is from the basic hair. There are all degrees of gradation between these two types. Also, well defined glandular hairs appear only in the more advanced members of the Malayan-Asiatic (Figs. 29, 30) and Afro-American (Figs. 14, 15, 19, 20) lines and hence correlate with other advanced traits. The Javan-Australian line has retained the intermediate basic to glandular hairs (Figs. 21-24). The origin of ray hairs is not so clear. In the Malayan—Asiatic line they seem to be derived from the basic type or one tending toward a glandular condition; there are all degrees of gradation between these types in P. grande and P. wandae (Figs. 27, 28). In the Afro-American line the ray hairs are often stouter, conical, and seem to have been derived from glandular hairs. These rays may appear as branches off a glandular hair in P. angolense, P. andinum (Figs. 14, 15), and occasionally in P. vassei, or may be independent but intermingled with glandular hairs in P. ellisi and P. vassei (Figs. 19, 20). Occasional hairs intermediate between the glandular and the ray condition may be found near the basal margin in P. vassei. The abundance of conical ray hairs also seems to indicate an intermediate condition between glandular and ray hairs. When the contents of the conical ray hair collapse, the ordinary ray with its flat shape is produced. Conical ray hairs are mostly found in the Afro-American line, and basic hairs are mostly absent from it. In P. ellisii, the ray hairs tend to originate from the center of the trichoblast cell, rather than from one side, as 1s frequently the case with hairs close to the basic type in origin. Nayar (1965, p. 13) has described hairs on Pyrrosia obovata which seem to be similar in many ways to the ray hairs of Platycertum ellisii. Pyrrosia heteractis, with the basic type of hair, is considered RHIZOME SCALES OF PLATYCERIUM 155 by Nayar primitive in relationship to Pyrrosia obovata. Nayar implied that the hairs which arise from the center of the cell and have dilated bases as in P. obovata are separate entities not related to protuberances in origin. It seems that hairs of this type in Platycerium are a result of cell reduction or failure of the basal developing hair cell to elongate properly. Similar dilated hair bases on glandular hairs may be found in some forms of P. vassei and in P. ridleyt. Unicellular to multicellular hairs—Most of the species have hairs with 2-4 cells. But one has only protuberances known as papillae (Fig. 25), others have mostly 1-celled hairs (Fig. 20), and others have mostly multicellular hairs (Fig. 26). It is generally believed that unicellular hairs are more primitive than multicel- lular hairs unless the unicellular condition is shown to be a reduc- tion (Netolitzky, cited in Carlquist, 1961, p. 34). In the Malayan— Asiatic line of the genus the primitive P. wallichii (Fig. 25) lacks hairs and has only papillae which presumably gave rise to 1-celled hairs. Hairs with more cells have evolved, and those of P. holttumai (Fig. 26) have 8 cells. The next species, P. wandae and P. grande (Figs. 27, 28), show a decrease in the number of cells to 5 or fewer. Basic hairs are still present, but semi-specialized hairs (i.e. inter- mediate glandular and ray types) have appeared. It seems that the appearance of semi-specialized to specialized hairs may be correlated with at least the partial loss of the basic hair types. In P. coronarium (Fig. 29), an advanced species, the cells in the hair are reduced to 2 and are glandular. The more advanced P. ridleyi (Fig. 30) has a large percentage of 1-celled glandular hairs. This reduction trend is also found in the Afro-American line, culminat- ing in P. ellisii (Fig. 20) which bears only 1-celled glandular or ray type hairs. It also seems that the advanced condition for spe- cialized hairs is from many to few cells. Incomplete stellate hairs—Ray cells are unicellular (Fig. 196) or are found as terminal or subterminal cells on basic or glandular hairs (Fig. 14). Two to several ray cells may appear on the apical cell of a basic or a glandular hair (Fig. 15); this condition ap- proaches the stellate hair. A less specialized development pro- igi ea US GD Te eS ie a Sa 156 AMERICAN FERN JOURNAL duces a long cell with many lateral ray-like cells along its axis; such hairs are occasionally found on fibrils. There seems to be a relationship between the stellate hairs found on the lamina and those on the rhizome scales. Rhizome scale hairs apparently show ontogenetic stages no longer present in the development of laminar stellate hairs. The development as studied by Straszewski (1915, p. 277) shows that a laminar stellate hair starts as a glandular cell which divides apically into successive glandular cells that elongate and flatten into rays. By studying a variety of hairs on P. angolense scales it is possible to reconstruct a plausible sequence of cell arrangement that could have led up to the stages described by Straszewski. One might take the point of view that the hairs on the rhizome scale represent reductions from the stellate condition. This seems unlikely, for the cells of a reduced hair would probably be com- prised of the component cells of the stellate hair type, Le. ray and glandular cells. Instead, a continuous array of basic to special- ized types of cells are present. Also, if reduction were the case, the most primitive species would have retained more of the stellate hair structure. This is not the case, however, for the species having this condition are not among the very primitive ones. The stellate hair tendency in scales is best developed in P. angolense and P. andinum. The other species have other special- izations or have not reached the stellate state. The separation of the ray and glandular cells into independent hairs is one type of specialization and is found in P. ellisii (Fig. 20). Flabelloid margin.—The cells of the scale margins are generally arranged longitudinally, but may be oriented at various angles, and may even be retrorse (Figs. 2, 28). Margins with cells that spread outwards in a fan-like manner Holttum (1957, p. 43) lescribed as flabelloid. Flabelloidly arranged cells may be absent, in clusters, or continuous, well developed or barely so, and they may merge with papillae and trichoblasts. If cells bearing protuberances, trichoblasts, and basic hairs are _ essentially similar and if they can unite laterally to each other in varying degrees, it is possible to explain the origin of the flabelloid RHIZOME SCALES OF PLATYCERIUM 157 margin by lateral fusion of ‘free’? marginal cells. This hypothesis is likely because there is no difference in color and texture between “free” marginal cells and those making up the flabelloid margin, the marginal cells being united in varying degrees up to a typical flabelloid state, and the orientation of the cells in the flabelloid margin being in the same direction as that of ‘‘free’’ hairs. That marginal cells as well as submarginal cells of a flabelloid margin may continue to produce papillae and hairs is further evidence that the cells of the flabelloid margin are basically the same as the marginal cells of unmodified scales. If the ‘‘free” hairs on a flabel- loid margin unite, a very broad, flabelloid margin as found in P. coronartum and P. ridleyi is formed (Fig. 2). The cells of these broad, flabelloid margins are oriented like many of the “free” marginal hairs in P. grande. In summary, the trend in the Malayan- Asiatic line seems to be from marginal cells arranged longitudi- nally to a narrow flabelloid margin with many ‘‘free” hairs to a wider flabelloid margin with few “free” hairs. This trend is absent or poorly developed in the other evolutionary lines of the genus. Where it does appear it is found in advanced species (P. vassez from Madagascar, P. bifurcatum, and P. hillii). False branching—False branching occurs in P. holttumai (Fig. 26) and occasionally in P. grande. It is characterized by having one or more hair cells with protuberances. Occasionally the pro- tuberances are separated from the cell by a wall, and thena truly branched condition exists. False branching may be inter- preted as a stage toward the fusion of hair cells to form a flabelloid margin. An alternative explanation is that these relatively newly derived basic hairs are still genetically close to the marginal cells and have retained their tendency to form papillae or, alternately, false branches. This seems to be the more adequate explanation, as many of the false branches are not distinct from the papillae of marginal cells. FUNCTION OF THE RHIZOME SCALES The most vulnerable part of these sometimes immense plants is the relatively small apical bud. Death of the bud results in death 158 AMERICAN FERN JOURNAL of the whole plant in species which do not produce lateral buds or buds from the roots. Rhizome scales function to protect the bud from desiccation, animals, and excessive water. Beneath the base fronds old rhizome scales may be found very much intact, covering and protecting the rhizome, although the softer parts of the scales may be worn away leaving only the sclerified stripes. Protection from desiccation is particularly important in these usually high epiphytes, as they are exposed to more drying condi- tions than low epiphytes or terrestrial plants. I do not know what animals might attack these plants in their native habitats, but the mass of base fronds is known to be a home for a multitude of organisms. Sowbugs, pillbugs, snails, and slugs are capable of eating through the scales and into the buds, particularly on young plants in cultivation. Small insects would probably have difficulty breaking through the scales covering the bud. I have observed scale insects unable to feed because they were entangled in the hairs of the rhizome scales. The insects were able to attach and feed where the rhizome scales ended on the phyllopodia. Smaller animals, however, could gain entry to the bud and rhizome through the injury inflicted by larger animals or microrganisms. The small larvae of fungus gnats (Mycelophilidae) apparently gain entry to the rhizome bud through leaf scars which are perhaps softened by decay or do not suberize well. Termites, birds, and various fungi also have been observed to make their home in the nest fronds as well. The base fronds usually form a ruffle of tissue over the bud, which diverts the flow of water. Although this ruffle may be said to serve other purposes, it also helps to keep the bud free of excess water. I believe that keeping excess water off the bud tissue is a very important function of the rhizome scales as well. Water- logged tissues are particularly vulnerable to microrganism infec- tions. If softened areas are produced from such infections, other organisms then readily find their way into the remaining tissue. The glandular hairs and large amounts of oil found in the rhizome seales of P. bifurcatum, P. grande (Joe, 1964, p. 71), and probably other species must also play some role in rhizome protection. RHIZOME SCALES OF PLATYCERIUM 159 Discussion With a knowledge of the rhizome scale characters described above and their distribution among the 17 species, it is possible to infer the kind of scales most likely possessed by the ancestral plant. In Table I, P. wallichii has the lowest value for specialized rhizome scale characters and probably has scales most closely resembling those of the ancestral plant. The relationships of present day species also may be clarified. For example, the general appearance of the South American species P. andinum could place it in any of the three evolutionary lines of the genus, but several rhizome scale characters and other less obvious structures place it closest to the African species. However, of broader interest are the changes in rhizome scale structures that have taken place along with the general evolution of the various species. Some of these changes are limited to a few species, but others are established patterns in this and probably other fern genera. The following changes which have taken place in rhizome scales of Platycerium seem to be correlated with evolution in the genus: fibrils present to absent; width narrow to broad; texture firm to thin papery, at least as a broad margin; apex filiform to non-fili- form and multicellular; base broadly attached to subpeltately to peltately attached; hairs restricted to the scale margin to spreading submarginally or centrally; margins mostly with papillae to mostly with basic hairs to mostly with typical rays and/or glandular hairs; hairs absent or basic hairs mostly 1-celled to basic hairs multicellular; simple glandular or ray hairs with many cells to the same with few cells; simple hairs to partly formed stellate hairs; and marginal cells oriented with the length of the scale to flabel- loid to very broad flabelloid. From the rhizome scale characters having phylogenetic value the following generalizations are made: 1. Elaborate scale structure is indicative of advanced species, except where reduction of the hair cells is involved.* 3 A similar conclusion is reached by R. M. Tryon, Jr., who finds that Cyathea seales with the least cellular ir Bt ea are primitive (in itt. Sept. 8, 1969). 160 AMERICAN FERN JOURNAL 2. Except in very primitive species, reduction in number of hair cells of simple but not stellate hairs is apparently correlated with increased hair specialization. 3. Hair development on some rhizome scales seems to reflect stellate hair ontogeny and shows stages now absent from the leaf. 4. The flabelloid margin seems to have developed from the union of papillae, trichoblast, and hairs. 5. Peltate scales developed by the folding under of the auriculate or rounded base and subsequent overtopping growth, rather than by the lateral union of the auricles. This is similar to the formation of peltate scales in Pyrrosia. 6. One of the important functions of the rhizome scale may be to protect the bud tissue from becoming water-logged. It seems that greater use may be made of rhizome scales in determining relationships within other genera, providing sufficient features are available. Scale interpretations will be more reliable with some prior knowledge of at least some of the relationships in the group. The use of scale characters will be of greatest help in interpretation of problematic taxa and in reinforcing or support- ing other phylogenetic interpretations. LITERATURE CITED Bower, F. O. 1923. The Ferns, vol. I. 359 pp. University Press, Cambridge Cartauist, 8. 1961. Comparative Plant Anatomy. 146 pp. Holt, Rinehart, and Winston, New York. Hourtum, R. E. 1957. The Scales of Cyatheaceae. Kew Bulletin 1: 41-45. Jor, Barpara. 1964. A Review of the Species of Platycerium (Polypodiaceae). aileya 12: 69-126. Nayar, B. K. 1961. Studies in Polypodiaceae, VII: Pyrrosia. J. Indian Bot. Soc. 40: 164-186. Nayar, B. K. and S. Cuanpra. 1965: Ferns of India-XV, Pyrrosia. Bull. Nat. Bot. Gard, Lucknow 117: 1-98. Srraszewsx!, H. R. 1915. Die Farngattung Platycerium. Flora 108: 271-310. WAGNER, as H., Jr. 1952. The Fern Genus Diellia, its Structure, reat mel Taxononiy. 167 pp. Univ. Calif. Publ. Bot. 26(1): 1-21 WAGNER, “W. H., Jr. 1964. The Evolutionary Pattern of Living Sls Bull. Torrey Bot. Club 21: 86-95. Los ANGELES City Coutece, Los ANGELES, CALIFORNIA 90029. SHORTER Notes 161 Shorter Notes STEM Cross-sECTION Prints Arp IN IpentTIFYING HoRSETAILs, —Stem cross-sections are often required for accurate identification of horsetails. These are difficult to obtain from dry and brittle herbarium specimens, I have found a method for making a printed record of the cross-section of a horsetail specimen quickly and with simple materials. It must be done before the specimen is pressed, but may be affixed to the herbarium sheet later as an accurate record of that aspect of the specimen and may be examined with the naked eye or under magnification. When a horsetail specimen is ready to go into the plant press, a sharp, transverse cut is made across the center of a mature internode. One of the cut surfaces is inked with a fountain or felt pen. Several imprints of the surface can then be made on smooth, non-absorbant paper (the field label if possible). Rough paper will make details difficult or impossible to see, and absorbent paper will change the print’s dimensions. Although the first prints may be inked too heavily, others will be perfect, and some will be too light. Unless a good labelling system is used to associate the prints with the particular specimens, the prints should be kept with the specimens in the press. The best prints should be pasted near the specimen label when the specimen itself is mounted.—Barpara J. GupMuNDSON, 5505 28th Avenue South, Minneapolis, Minn. 55417. PsILoTuM NUDUM SPREADING NorTHwarp.—In 1942 the “Whisk Fern,”’ Psilotum nudum, was recorded in Brown and Correll’s “Ferns and Fern Allies of Louisiana” as having been collected growing wild in a greenhouse, Krack’s Nursery, north of New Orleans, having apparently been accidentally imported with potting soil. A recent publication (Sida 3: 525. 1970) by Dr. G. Rhodes now reports the species as growing wild in four Louisiana Parishes—Terrebonne, St. Mary, Vermilion, and Lincoln. In the last named the species is some 250 miles north of any previously known locality. It is thus apparent that this plant is spreading northwards rapidly, perhaps from wild plants occurring to the eastward in Alabama or Mississippi.—C.V.M. 162 AMERICAN FERN JOURNAL TERMINOLOGY OF THE SPORANGIAL STRUCTURES OF EQUISETUM. —Several years ago a population of Hquisetum X litorale with many abnormal cones was discovered by Dr. W. H. Wagner, Jr., near Milan, Michigan. The population was extensive, and had invaded an old bean field. In this dry, sandy habitat, atypical for this hybrid, the strobiliferous stems frequently had cones with proliferated tips (Fig. 1). The apex of the cone, instead of ceasing growth as it usually does, reverted to a vegetative condition. This type of teratology has been previously recorded in various species of Equisetum, and Gluck,! Kashyap,” and Tschudy* have described the intergradation of fertile and sterile appendages associated with the transition from the strobiliferous to the vegetative apex. The same intergradation, seen here in E. X litorale (Figs. 2-7), reveals an innate capability of the appendages produced by the apex to develop either as vegetative leaves or as sporangium-bearing structures. This indicates that the two structures are homologous, and that the term ‘“‘sporophyll,” a more exact term than “spo- rangiophore,”’ should be used for these structures. Tschudy* argued for the term ‘‘sporophyll” on the basis of teratology in EL. telmatera, and I wish to reiterate it here on the basis of teratology in LE. X letorale. Since the leaves of Lycophytina, Sphenophytina, and Pterophy- tina are all apparently a) pendently evolved, calling their sporangium-bearing structures “‘sporophylls” does not imply any homology between the sporophylls of these groups, but only be- tween the leaves and sporangium-bearing structures within each group.—Richard L. Hauke, DEPARTMENT OF BoTANY, UNIVERSITY oF RHODE IsLAND, Kineston, R. I. 02881. 1 Die Spor ophyllmetamorphose. Flora 80: 303-387. 1895. 2 Some abnormal cones in Equisetum debile. J. Indian Bot. Soc. 9: 240-241. 2 The significance of certain abnormalities in Equisetum. Amer, J. Bot. 26: 744-749. 1939. AMERICAN FERN JOURNAL VotumE 60, PLate 20 Fig. 1. Cones or E. X LITORALE SHOWING PROLIFERATION OF THE TIPS. Figs. 2-7. SERIES OF SELECTED APPENDAGES FROM THE CONES IN Fi. 1, FROM A TYPICAL SPOROPHYLL TO A NEARLY TYPICAL SHEATH SEGMENT (LEAF), 164 AMERICAN FERN JOURNAL Kinps or CYsToPTeris IN CALIFORNIA.—Recently John Thomas Howell, while preparing a floristic account of the ferns of the Sierra Nevada, suggested to me a study of the two spore types of Brittle Bladder Fern in that area. I enlarged the project’s scope to include all the California collections of Cystopteris in the Cali- fornia Academy of Sciences herbarium, more than 120 spore- bearing collections. Based on spore characteristics, two separate taxa are recogniz- able, although in general appearance they look identical. Profumo’ examined the gametophytes of the two types and also found no morphological differences. At times, the two types have been considered separate species, C. fragilis (L.) Bernh., with spiny spores, and C. dickieana Sim, with non-spiny (rugose-verrucose) spores; the two have also been considered subspecies. The specimens I examined show that both kinds may occur in the same locality, a fact which was recognized by Profumo and by Hagenah,? who occasionally found both mounted on the same herbarium sheet. Within the Sierra Nevada, 14 collections were the fragilis type and 67 the dickieana type, a noteworthy excess of the latter. Non- Sierran collections showed 22 fragilis type and 20 dickieana type, a more even distribution. While both kinds occasionally may occur together, there appears to be a predominance of the dickieana type at higher elevations. This corresponds to the findings of Larsen’ in Greenland, who suggested that C. dickieana was a high Arctic species and that C. fragilis was a sub-Arctic species. Perhaps these two taxa should be considered as forms rather than subspecies since their distributions are essentially identical and the only apparent difference is in their spores—Rospert B Setzer, Allan Hancock Foundation, Dept. of Biology, Univ. of Southern California, Los Angeles, California 90007. dic fa = a. Webbie 9. The gametophyte of Cystopteris fragilis and Cystopteris * Hagenah, D. J. 1961. Spore studies in the genus Cystopteris. I. The ee. bution of of ‘Ceseeaes with non-spiny spores in North America. Rhodor: 3 Larsen, K. 1952. Lasaasne sae i Coco ag oo fragilis coll. med piggede og vortede sporer. Bot. Tidsskr Norres AND NEws 165 Notes and News A New Newstettrer.—To better promote communication and the exchange of materials between our members, the American Fern Society is inaugurating a quarterly Newsletter. Prof. James Montgomery, an enthusiastic student of North American ferns and a frequent participant in our Fern Forays, and Mr. F. Gordon Foster, an experienced horticulturist and author of “The Garden- er’s Fern Book,” will be in charge of the Newsletter. Non-technical items and notes of passing interest will be most welcome. The officer’s reports and a membership list will appear in the News- letter each year. We hope especially that our members who are naturalists and gardeners will contribute to the Newsletter and will find it interesting and informative. GrEoRGE NEVILLE JongEs, Professor of Botany and Curator of the Herbarium at the University of Illinois, died last June 25. He had been a Fern Society member for a quarter of a century. He was the author of many monographs and floras, including “A Botanical Survey of the Olympic Peninsula” and “Flora of Illi- nois.”” His principal work on ferns, “‘An Annotated Bibliography of Mexican Ferns,” was published four years ago. PLANTS OF ASPLENIUM EBENOIDES X CAMPTOSORUS RHIZOPHYL- LUs are available to persons interested in growing this backcross hybrid Spleenwort. Please request living specimens. Scott’s Spleenwort, A. X ebenoides, is the hybrid of the Ebony Spleenwort, A. platyneuron, and the Walking Fern, Camptosorus. We have available numerous proliferations of the backcross of the hybrid to the walking-fern, which was originally made by Mrs. Kathryn E. Boydston at Fernwood, Niles, Michigan. This backcross plant has proved to be extremely vigorous in culture. It looks like a some- what lobed Walking Fern with a dark stipe, and it grows in a —— spreading manner. It should be grown in a Wardian case or terrarium in ordinary potting soil—W. H. Wacner, JR., M pia Botanical Gardens, University of Michigan, Ann Arbor, Mich. 48104 166 AMERICAN FERN JOURNAL -Epwarp M. Suieips, 1895-1970—Members of the American Fern Society, especially those who some years ago participated in its field trip in eastern New York, will regret to learn of the death from a heart attack on September 17th of ‘“‘Eddie” Shields, who on the occasion of the field trip entertained us at the summer residence he then maintained at Haines Falls in the Catskills. As a youth Eddie roamed his native Chester County, Pennsyl- vania, collecting at first minerals and Indian relics. Receiving from the family real estate holdings there sufficient income to live comfortably, he became an amateur naturalist of wide interests. On a field trip in Florida he made some notable orchid finds, thereby becoming acquainted with Oakes Ames, the specialist on these plants. His interest in ferns was aroused when, on a visit to Cambridge, Ames introduced him to Charles A. Weatherby. Exploring the Haines Falls region for ferns, he discovered a remark- able southern disjunct outlier of the Fragrant Fern, Dryopteris fragrans. ae After some years, Eddie came to spend his whole time in Sara- sota, Florida. Studying subtropical horticulture, he developed around his house a garden including many uncommonly used yet highly attractive plants. In an adjoining slat-house, which after a couple of disastrous freezes evolved into a greenhouse, he special- ized on growing unusual orchids. He would stop in at one of the many Florida orchid-sales places, and with his keen eye and thor- ough acquaintance with current literature, would often detect as ararity a plant offered as a common one. Ferns in his garden greenhouse were merely incidental; in ake living room he had a spectacular hanging-basket culture of Narrow Strap-fern (Campyloneuron angustifolium). His fern books and herbarium were given to the West Chester, Pennsylvania, State College. _ Eddie is survived by his wife “Jerry” and by several children and stepchildren. Memories of his cordiality and generosity to me over the years have made this note most difficult for me to write. —Epear T. Wuerry, Leidy Laboratory, University of Pennsylvania, Philadelphia, Pa. 19104. REcENT FERN LITERATURE 167 Recent Fern Literature Paciric NorTHWEsT FERNS AND THEIR ALLIES, by T. M. C. ft aheee eae of Toronto Press, 33 East Tupper St., Buffalo, N. Y. 14203. 1970. 247 pp., illustr. $15.00.—This is the first full- a nee fern flora to appear in several years. The area covered includes that treated recently in the ‘Vascular Plants of the Pacific Northwest” (reviewed in this JouRNAL. 60: 34-38. 1970), but is more extensive, since all of British Columbia, the Yukon Territory, and all of Alaska is included. The number of species treated is 97, about a quarter of all those known from North America north of Mexico. Many of these are boreal plants familiar in the eastern United States and Canada and many occur also in Eurasia. Twenty-seven are western American endemics (but not local endemics, except for Botrychium pumicola, confined to the region of Crater Lake, Oregon). The species all have full descrip- tions and most of them have distribution maps and line drawings. The latter are very good, so good that most species can be easily recognized from the drawings alone. (I would except the drawing of Polystichum scopulinum, which shows a plant not typical of this species and which might even represent P. kruckebergii instead.) The systematic treatment is commendably conservative. The comments indicate the variations found and some of the taxonomic problems still unsolved. There are few errors indeed, and the comments below are of strictly minor importance. The authority of Blechnum spicant is not “J. Sm.” but Roth (cf. this Journal 34: 51. 1944); incidentally, it is usual to use “‘J. Sm.” for John Smith; the author intended here was not John Smith but Sir James Edward Smith, usually cited as J. E. Smith, for the abbreviation of Smith to “Sm.” is really not sanctioned by the Code. The work in which Dryopteris was published by Adanson is not ‘“‘Farn Pl.” but ‘Fam. Pl.” In the description, the sori of Polypodium montense are said to be “‘submarginal,” but in the drawing they are shown as about medial (for a comment on the name ‘‘montense’” see this JouRNAL 60: 126-7. 1970). Polypedium virginianum is described as having the segments “‘linear-oblong to 168 AMERICAN FERN JOURNAL lanceolate ... obtuse at apex,’? which is essentially correct, al- though the apex does vary from rounded to acute, but in the com- ments it is said to be distinguishable from P. montense by the segments being ‘‘narrowly ovate with acute tips,” which is not true and which contradicts not only the description but also the draw- ing, which shows the segments as oblong and rounded at apex. In the synonymy of Thelypteris nevadensis, the first line ought really to be ‘‘ex Morton, Amer. Fern J. 48: 139. 1958,” as required by the Code; otherwise it appears that Clute published the name some eight years after his death. The A olla of the area is referred to A. mexicana, following Svenson; the report of A. filiculoides ai Alaska by Svenson is Reece with the statement “almost certainly an aquarium escape,” which is perhaps an unwarranted assumption, since the specimen was collected in 1868, an early period in the history of Alaska when aquaria must have been essentially unknown; still, the record must be considered doubtful since no other Alaskan material has ever turned up. There are specimens in the U. 8S. National Herbarium from Oregon and Washington that have been identified as A. filiculoides (and sub- sequently to Svenson’s treatment), but they are sterile, I believe, and so somewhat doubtful. An effort should be made to collect fertile material from these states. Possibly Svenson overstressed the character of the septae in the glochidia, and his treatment has never been critically evaluated. Although it is irrelevant here, I might mention that although Svenson indicated that A. caroliniana must spread solely by vegetative means, since megaspores are never found in American material, some recent European workers have described the megaspores of this species, considered intro- duced in Europe. It is possible that the European plants so identi- fied are not really A: caroliniana, and the matter should be in- seventies. It is rather too bad that this book perpetuates the llings ‘‘sabinaefolium,” ‘‘matricariaefolium,” and ‘‘andromedae- folia”: although these were the original spellings, Art. 73, Note 2 says that the use of a wrong connecting vowel is to be treated as an orthographic error, and Art. 73 itself indicates that orthographic errors are to be corrected; thus the correct spellings are “‘sabini- AMERICAN FERN Socigery 169 folium,” “‘matricariifolium,” and ‘‘andromedifolia.”’ One feature, which does not at all detract from the book but which is sufficiently peculiar to deserve mention, is the placing of Selaginella at the end, following the Polypodiaceae, instead of at the beginning, with Lycopodium, Isoétes, Equisetum, and other fern allies —C.V.M. American Fern Society New Members Mrs. Rhoda D. Bancroft, 4 Willard Ct., Norwich, NY 13815 Mrs. Henry W. Bookout, Jr., 120 Smith St., Islip, NY 11751 Mr. Parul Catling, 104 Victoria Park Ave., Toronto 13, Ont., Canada Mrs. Thomas A. Cole, 1000 S. Grant Ave. . Crawfordsville, IN 47 933 Mr. Edward L. Davis, Dept. of Botany, Buiy: ot Massachusetts, Amherst MA 01002 Mr. Robert B. Faden, East African Herbarium, P. O. Box 5166, Nairobi, Kenya, Africa Mrs. Leslie J. Helmstetter, 2546 Hillsman St., Falls Church, VA 22043 Mrs. J. J. Howell, 4028 Greenwood Dr., Ft. Pierce, FL 37450 Mrs. 8. H. Inglis, 233 Red Fox Road, Stamford, CT 06903 Mr. R. Kean Ivey, Jr., Box 108, Rt. 3, Charlottesville, VA 22901 Mr. Terry W. Lucansky, 3600 Tremont Dr., Apt. H-3, Durham, NC 27705 Mr. Donald McFall, 4711 Forest Hills Rd., Rockford, IL 61111 Mrs. Robert D. Moody, 7705 Lynch Rd. ’ Sebastopol, CA 95472 Mr. Richard Moyroud, 202 Grove Way, Delray Beach, FL 3 versal Mrs. Jane W. Pettus, 805 South Warson Rd., St. Louis; MO 6312 Mr. Michael G. Prive, Dept. of Botany, UPCA, College, Laguna, phe Mr. Arthur Raguse, 715 Linwood St., Abington, MA 02351 Rev. Pau : John Rich, First Parish Unitarian Church, E. Bridgewater, MA Mrs. Wiuse, J. Wright, 1914 W. Division St., Arlington, TX 76010 Changes of Address Mr. Michael I. Cousens, Dept. of Botany, Washington State Univ., Pullman, 99163 ee Mrs. V. De Benedictus, Zoology Dept., Syracuse University, Syracuse, NY 13210 Dr. Delzie Demaree, 109 South Ave., Hot Springs, AR 719 Mr. William es Fulcher, Biology Dept., Guilford sci Greesnboro, NC 2741 170 AMERICAN FERN JOURNAL Index to Volume 60 Abacopteris, 120 Aberrant Leaves on Angle-Shoots of Selagi- nella iastines Soothe, Actiniopteris radiata, 137, 138 Acrostichum alcicorne, 7-11; bifurcatum, 9; as stemaria, 7, Adiantaceae, Adiantum, 124, 137, Se ee ee. 138; capillus eris, 128, 138; caudatum, > 138; pedat aa 138; lunulatum, Bee 8 ium, ae spinulosum, 24; thelypteris, 24 Aspidotis, 1253 mee 40 adiantu: im-nigrum imum, oli platyneuron, 131, 133; ruta- septentrionale, 129-133; sylvaticum, 122; trichomanes, oe 139; tripteropus, 86; Varians, 1 vestitum, 122 sai bao ‘ Jfilix-femina, 39, eV ‘Cory ict gee 100; ifer lilloi 103; avpcaniter | pate re S oat aes 168; filiculoides, 168; Asplincean, 2 Bajpai, Ni eee (see B. K. Nayar) Pelt an, Frances (see L. Lee) vee) at phy to Floras of Southeast Asia rev.), 5S. A Note on the ane of ‘Cvsiomecia bss eg Blechnopsis malac ea 1a Blechnum, 104; 105; australe, 104; “oodioides, "36; gr num, EN a a indicum, 121; 105 1 sie Wg KA 9 35, 167; tabulare var. setigeru’ 139; lunaria, 139; | esi ; Cardillo, Frances M. Stelar Anatomy of Six Species of Lycopodium, 89 Caeues ot perce cg ge Chandra, Let Pfc we commen egy of the Rhizome of Cibotium baroi z, 68 Cheilanth rig 125; catanensis, 128; fragrans, 128; gra cillima, 40; lanosa, 36; siliquosa, _ Chromosomes and Speciation in Ferns (rev.), Cibotium, 68; Reha 68, 70, 71, 130; Cornopteris, 98, 100 Cousens, M, I. £4. T: Sead deeeretge cick Ontoge eny and Sex re, xpres Dry pteris Indovieiana, 1 wie So the ann ei Hitchcock) togra peridaea Cyathea, 38, 71, 159 Cyatheaceae, 68, 71 pike princeps, 118 y closi 126; Saert ae s, 1 : Cystopteris, 98, 100, 164; subg, A opteris, bulbifera, 98; subg. Cystopteris, 3 ‘Ste hana 100; dickieana, 164; PIA ce 98, 100: fragilis, 98, 100, 164; montana, 35; tenuisecta, 8-100 Uae segrag gat 33, 39 DeVol, el he age of Taiwan-5. Hym Pac Bie richomanes (rev.), 87 Distaneynceria, cd aes pega magellanicum, 106 Diellia, Di Sie sineela vanica, 100 Diplazium, 38; feces 98; grammitoides, 122; sylvaticum, 122; tenerum, 122; vestitum Dipteris, 125; conjugata, 125 1 P 167; assimilis, 38; aus- triaca, 37, aust: rome 37; barbigera, enn ‘earthusiana, Boy hrysoco 137; Bee 5s ate, 38, ppideteneg 34, 25, 37, 39 intermedia, 83; ludo- Vicia ana, go 21, ap om 26, Soe, ata, 137; odontoloma 137; schim periana, 3% i; spinulosa, Of. "33, att villastl m, 73, 74, {howe 125; carly nudatum, 103; - Ronnie um, 75; erinaceum, 75; eximium, ashe inieri, 75; hirtum, 75; MS jerome ity ee, 76; longifolium , 75, vens, 75, Fie wiaexincn, ‘75: tectum, ‘75, WW; undulatum, . W. Two New State Records for Majo rth American Range Extension for tne F ay Spleen- aoa um ceptentrionale, 12 um, 162, 169; arvense, 137, debi 140; x Gerais, 162, 163 um, 128; te eeenags ~~ 162 whe Fern Collec Some European Herbaria os Fern Hisbridiatne at the University of Leeds Botrychium vir- Attac hed to lege ee ein 62 rine Ps Pal péskion, Pa A Further Note on the oat of Platycerium alcicorne, 7 Gametophyte if ae aed boat Expression n Drypoteris ludovi Gleichent 28 Goniophiebiam, 66; By ean 67 Goniopteris tetrago' Cocmamaiddiene 123 Grammitis, 123; albidula, 66; crispata, 66; sect. Cryptosorus, 66; discolor, 66; eminens, INDEX TO VOLUME 60 66; kalbreyeri, 66; mathewsii, 66, 67; sechellarum, 124; trifurcata, 66 Gud = undson, Barbara J. Stem Limiting Factor in the Tsteibution of Seobten. epee oka e 134 Hauke, n, S. G. (see H. A. Hyde au . Term rminology of the Sporangial Structures ‘of Equise 162 Hel minthosta: ” ys zevlanica, 140 Hitcheock, C. A. Cronquist, M. Ownbey, &J.W.T Puatiane pvascular bhue ry of the Pacific he est, Part 1 x" Holttum pei = mee mentary he sptimd oO Type on barium of KB B. Pres Pie u 119; A Rata Flora of Malaya, vol. II—Ferns of Malaya, ed. 2 (rev)., Horner ao il I. I. Cousens) Hoshizak i, Ba A ti ce The Rhizome Scales o Hymenophy vm 06: ellanicum, 105, Pe td eselifoliu: Pat al Vormpire as 107, 114-117; crenatum, 98, 100. 1 107, 1 112, 115 The Thadidated Flora of Illinois: Ferns (rev.), —— Pteridophytes Used in Folk Remedies, Tsoétes, 30, 35, 169; sect. Ee oe Si; echinospora, on 134, 335; Bold braunii, 37, 135, ie engelmannii, 0; melan opoda, res be setacea, 33. sect. Tuber- cults ae, 3 ug Fal 37 Karyologic al Studies on Aspleniaceae 1. Karyotypes of Three Species in Asplenium LY (re Kawakami S. (see S. Tatuno) Kinds of Cy stopteris in California, 164 — rthwestern North ey Lectotype of Polypodium leptophyllum Ate me ” Frances Belknap. Hard Water as a Limiting Factor in the Distribution of 34 Ospora. indsaea, 124 Lloyd, R. M. A Survey of Some Morpholo ay cal Features of the Genus Elaphostemenn n Costa a, 73 ana, 104; ger 104, 105; 9 ‘lan ion var. angus Getta: "105; 0; agai 103, 104; setigera, . Fern Hybridizing at the Uni- of Leeds (rev.), 86 9 ; lue’ ri tryoni nii, 5S: Ghacaun: 91, 92,95; | poro on igheogs 33, 131; selago, 141; tristachyum, 171 Lygodium, 38; circinnatum, 141; flexuosum, 141; japonicum, — microphyllum, 141 A Major North American Range Extension tue Forked Cotes mwort, Asplenium prt fo le, Marsilea, 35; minuta, 128; quadrifolia, 141 Milne magellanic a, 106; quadripartita, ne oe fos, 126 | = aay = icone: and Speciation in Kohtiaacek: i. A, igar” Fe stersper Flora of eee e. Ferns iat Further ee n the Type prea Pyrro rinceps, a Fern Nee teC ulti, vation, “iis. Taxonomic Notes on Ferns, 103 Nayar, c < Nis Rei Vestigation o f the Mercieee re Hy ma ematium crenatum, 107 Nephrolepis, 116 Neu wpe rade ‘on, 7 A New of Micro pia speluncae, 28 A New Cane: for a ten of Polypodium from Northwestern North America (rev.), ee costatus, 121 A Note on the Gametophytes of Cystopteris tenuis: Notslacsn: a, 125; ae 128 Ophi — 60; lusitanicum, 128; vulgatum, Olenhs. 81 Osmunda regalis, 1 Ownbey, M. Asi c. 1 Witakosk : Pacific Northwest Ferns and Their Allies re: rev.), Bacay Species of Grammitis, 65 Phayontente connectilis, 131 Phillips, W. S. Psilotum nudum New Arizona, Bhs sagittata, 128 425, - 126; to 120; rium, 121, bie red 159; aethiop- sf alci rne, ee andinum, 141- 146, 148, 153, 154, 156, 1 lense, ae. fare vest 11 146, banerjiana, pe _— goo 151, > sk ie 144, 146. 9-155; willin Fyoee, 149, 151, 160; heteractis, 154; obova , 154, 155 172 Pleopeltis, 125 Poveda 34, 106, 123 Polypodium, 80, 124136: albidulum, 66; eur 24; alternifolium, 120, 27; australe, 85; cam- brides ry soutetak. 12]; discolor, 67; hesperium, 35, td heterophyllum, 102; ri, ophyllum, 101- 103; rip: sechellarum, 124° ieliarsataan, 127; var ium, 102; virginianum, 167; vulgare, 85, 142, var. hesperium » 35, var, serratum, 128 Pulyatiohian, 35, 116, 124; iruckebereii, 167; scopulinum, 167; squarrosum nad 120° Profe Pron 120 Psilotum 1, 32° nu udum, i a, 30 Psilotum nudum Sooeeding Northward, 161 Pteridium, 25; ie 43, 25, , 142 The Pte ridoph: of Teiwann-6° “aon no- phyllaceae: Trichornanes (re vd, Pteris ensiformis, 142; multifida, 142; quad- riaurita, 125; serrulat ta, 142; vittata, 128; wallichiana, i ito Ptychophyllum magellanicum, 1 Puri 8. Indian Peaciepb sie Used in Toe Remed edies, 137 Pyrrosia, 118; costata, 119, 121; grandissima, 119; lingua princ eps, 118, 119; splendens, 1 Pyrrosia princeps, a Fern New to Cultivation, 118 Reed, Clyde F. ri nye ag to Floras of Gootiaah Asia (rev.), 39 A Reinvestigation ge, Frei ey of Hypodematium crena ee Report of: Auditing roe 45; Fern Spore Exchange, 46; Judge: of Traihicna; 45: oh ce t, 40; — pals Treasurer, 42 oat s of Tropical itoaa Pterido phyta, Epi ae Bibliography to Floras of Southeast penal 8 bag romosomes and Speciation in erns, . oo eninge ry on ‘Some T ype Sectnens ar in the Herbarium of K. B. Presl, 19; Fin ybridizing at the University of Leeds, 86; toc Palaestina, Part One, Text, 127; The soley Flora leg pecoetnd Ferns. ems, 32; K Kary: i ree Species: in Raleeiuk: 86; A utp a a of a from oe es AMERICAN FERN JOURNAL hg of Malaya, vol. Il—Ferns of Malay: ed. 38; Vascular Plants ie: the Pacific Meniwek part 1, 34; The rian Fern 84; We Ish Ferns, Cl aliases Quillworts, and Horsetails, A Descriptive dbook A Revised Flora fron ie vol, Ii—Ferns aor Malaya, ed. 2 (rev Sali nia uricula raniitalin. 124, 125 Salviniaceae, 123 Sch Pots e, hat A. GC) Ls. riews of Tropical Afri eridophyta, i phos 2 123 Senos @ ag 142, 169; martensii, 1-3, 5, 6; Digge ae'9g 2 ,5 Setie K inds of Cystopteris in Cali- ‘er Peay oe of Six Species of Lycopodium, ie ees a Prints Aid in Identifying go s, 161 Stenochiaena usta, 15 123; e Morphological Features capheoides um in Costa Rion: 13 hart r % "Karvological St em on Aspleni aryot hree Species in tein (rev stg 86. Tosconsae } Notes on Ferns, IV, 103 Taylor, T. M. Pacific Northwest Ferns and Their Allies (rev.), Tectaria, 34, 116; polymorpha, 142 Receninolegs of the Sporangial Structures laurifolia, 122, quisetum, 16 be fooslaget 24, 34, 124; dentata, 13, 14, laph yropteris, 103; megapili, Trichomanes, 87, 124; scandens, 8 Two New State ‘Records for Isoétes in Ken- tucky Vasbalat Organization of the Rhizome of Cibotium barometz, 68 Vase — * ig of the Pacific Northwest, Part The Victorian, Fern Craze (rev.), 84 Vi ae sie eae Welsh Ferns, Clubmosses, Quillworts, and Horsetails, A Descriptive Handbook (rev. b Ww ia, 33, 34; ilvensis, 34; obtusa, 34 Ripon radi chamissoi, 36; fimebriate, 36; 36; radicans, 36 124; erica, 126; Sta Pais Their Allies, 167; gone terid: Rinhdpseci. we 124, albo brunnea, phyta of Taiwan—-5. Hymenophyllaceae villosissima subsp. subpinnata, 124 Trichomanes, 87; Reviews of Tropical Zohary, M. Flora Palaestina, Pees One, African Pteridophyta—1, 123; A Revised Text (rev.), 127 Errata Page 105, line 3: For “‘B. araucana” read “B. araucanum.” Page 125, line 8: For “A. auriculata’”’ read “‘S. auriculata.”’ ” ‘Page 127, line 8: For 1166” read “11667. TRIARCH INCORPORATED =A Service to Biologists = To better serve the biological community: 1. TRIARCH produces and stocks nearly 3000 dif- ferent prepared microscope slide items for use in gen- eral botany, plant anatomy, plant physiology, phyto- pathology, bacteriology, general zoology, vertebrate histology, vertebrate embryology, and parasitology. 2. TRIARCH offers to prepare slides of new or spe- cial items for a nominal fee or in exchange for pre- served materials. 3. TRIARCH advertises in biological journals to help support the financial needs of the associated So- ciety. For our current catalog No. 15, or to request special service, write to: Paul L. Conant, President TRIARCH INCORPORATED | Ripon, Wiseonsin 54971 American Fern Journal A QUARTERLY DEVOTED TO FERNS Published by the AMERICAN FERN SOCIETY e Eprrors DAVID B. LELLINGER C. V. MORTON ROLLA M. TRYON IRA L. WIGGINS VOLUME 61 i ee ae MONUMENTAL PRINTING COMPANY, BALTIMORE, MARYLAND Contents Vorume 61, NumpBer 1, Paces 1-48, Issuep Marcu 23, 1971 A Study of Azolla pinnata R. Brown........ A. Sweet anp L, V. HILus A New Variety of Lindsaea odorata Roxb. from aie icici in the WN SiS pase en | Set ue Ch tals ee T. Sen U. Sen The Proper Disposition of Meniscium macrophyllum Kunze C. V. Morton The Thelypteris normalis Complex in the Southeastern United States A. Rei Smita Asplenium pinnatifidum x trichomanes—A New Record for Indiana ERALD J. GASTONY mas WyWart Ostricke oRern 360 ct 4 erates eb eee H. Lou Gipson Shorter Notes: Niphidium longifolium, a Necessary New Combination; Bulbous Adder’s-tongue Common in Louisiana...............--.-- mecent- Perm: Literature. oa oe to ee ees ce Shei American Fern Society : Report on the 1970 Fern Foray............-..-- Votume 61, Number 2, Paces 49-96, Issuep June 24, 1971 Lindsaea (Schizoloma) ensifolia Swartz in Hawaii.W. H. WaGner, Jr. The Fern Collections in Some European Herbaria, VII C. V. Morton An Appendageless Psilotum. Introduction to Aerial Shoot Morphology ERT S. ROUFFA Vascularization of Fern Leaves Rosert C. LomMasson anp C. H. Youne, JR. Shorter Note: A Wrongly Localized Species of Pyrrosia..........---. Recent Fern Literature Votume 61, Numer 3, Pagsas 97-144, Issuep Novemser 15, 1971 The Fern Vegetation of Aldabra Atoll................. F. R. Fossuerae Leaf Epidermal Studies in Marsilea Joun T. Micke ANb FRANK V. Vorava — c=} Pa 101 The American Species of Plagiogyria sect. Carinatae Davin B. LELiincer The Genus Stenochlaena J. Smith with Description of a New Species R. E. Hotrrum The Variation in Spore Size and Germination in Dryopteris Taxa Dean P. WuittierR AND W. H. WaGNneR, JR. The Gametophytes of Natural Hybrids in the Fern Genus Pellaea Tuomas R. Pray Schemochromic Blue Leaf-surfaces of Selaginella Denis L. Fox anp James R. WELLS Shorter Notes: Asplenium kamtchatkanum Gilbert, a Dubious Eastern As‘atic Fern; A New Locality for Psilotum nudum in Sonora, Mexico. . MONT WET TUMOR ER: ios es oh ha ov ch da Sasa oe ee ane VoLuME 61, NuMBER 4, Paces 145-192, Issuep DecemMpBeEr 17, 1971 CONTENTS The Genus Grammatosorus. .........5.0c0ccecceeeces C. V. Morton The Genus Selaginella in Baja California, Mexico...... Ira L. Wiaeins Notes on the Ferns of Dominica and St. Vincent, II C. V. Morton aNnpb Davin B. LELLINGER Two New Tree Ferns from Costa Rica............ Luis Disco GémeEz Comparative Habitat Requirements for Spore Germination and Pro- thallial Growth of Three Ferns in Southeastern Michigan Royce H. Hiti The Gametophyte of Thelypteris erubescens. ... LENETTE R. ATKINSON Shorter Notes: A Recent Find of Isoétes in Louisiana; Jamesonia alstonii in Oaxaca, Mexico. ¢: 2556 ee oe 6 ie ss Recent Pern Literature 63 oe a ee ete Index to Volume Gio. oe a ee he ee ae ee Vou. 71 Cos4 January-Marcsa, 1971 No. 1 American Fern Journal A QUARTERLY DEVOTED TO FERNS Published by the AMERICAN FERN SOCIETY Cc. V. MORTON ROLLA M, TBYON A Study of Azolla pinnata R. Brown........A. Sweet anp L. V. Huis 1 A New Variety of Lindsaea odorata Roxb. from Darjeeling in the PUN on ak Ee ck T. Sen anp U. Sen 14 The Proper Disposition of Meniscium macrophyllum Kunze C. ¥. Morton 17 The Thelypteris normalis Complex in the Southeastern United States A. Rem Suita 21 Asplenium pinnatifidum x trichomanes—A New Record for Indiana Geratp J. Gastony 32 See Atha’ aeeens Sonaeie ae elle ee ag GOT 37 pau American Fern eaciele ae oat Council for 1971 Warren H. Wacner, JR., > peau of Botany, University of Michigan, Ann Arbor, Michigan 48104. President Joun T. Mucxer, New York Botanical Garden, Bronx Park, Bronx, New ‘York. 10458. Vice-President cnaRp L. Hauxe, Department of Botany, University of Rhode Island, Kingston, Rhode Island 0288 Secretary LeRoy K. Henry, Division of Plants, Carnegie Museum, Pittsburgh, Penn- sylvania 1521 3. Tre easu: Davw B. Letuinaer, Smithsonian’ Institution, Washington, D. C. 20560. Editor- in-Chief - Pational Society Representatives Warren H. WacNer, aa A.A.A.S. Representative Routia M. Trron, JR., Harvard Uni A.1.B.S. Representative American Pe Journal EDITORS Davi B. LELLINGER..... Smithsonian Institution, Washington, D. C. 20560. “3 V. Morton...........Smithsonian Institution, Washington, D. C. 20560. ray Herbarium, Harvard University, Cambridge, Mass. 02138. Tra L. 4 sansa de ..- Dudley og hima Stanford University, SStanford, Calif. An illustrated quarterly Snes to the general vag Rt ferns, owned by the an Fern Society, and published at 3110 Baltimore, Md. 31211. fectakdins = Pen paid at Baltimore. The pages ges of the Jour- —— open to members who wish to exchanges; membership lists arrange ex interested in obtaining specimens from different Se ag for publication should be addressed to the . pe ee al ig aa back numbers o {64 pages or 00 each; over 80 pages, $2.50 cts Cumulative lr to Volumes 15,3 1-25, 50 cents. Ten ‘en percent discount on orders of six volumes Librarp and Berbarium Dr. W. H. Wagner, a Departmen ment of ney eid , University of t Michigan, books and specimens at any time, the ianunee Goa teic alt videghas come, lebs, $1.25 ++ M Neill D. Hall, Ne 2 1225 ae tb 95th Street, pocerapel Washing 98115, is Director. staat reee cdg bee age ee and Bequests | dis sal indents ts ha eae estes ati its services to mem- Se these ap cto oe gh ese seo pen Se pe American Fern Journal Vot~7i C$) A JANuARY-Manrcn, 1971 No. 1 A Study of Azolla pinnata R. Brown A. SwEET AND L. V. Hits! A study was undertaken to ascertain the morphologic variability of the vegetative and fruiting structures of plants within the Azolla pinnata complex. Plants of this group have been commonly referred to three species, A. pinnata, A. africana, and A. imbricata. Our observations indicate that only one species is present, although two varieties based on vegetative morphology can be established. The synonymy given by Christensen (1905, p. 148) forms the basis of our synonymy for A. pinnata. In addition to Christensen, the original diagnoses and the work of Baker (1887) and Shen (1961) were consulted in working out the taxonomy of this group. Anatomical studies have been done on these plants by Griffth (1845), Strasburger (1873), Sud (1934), Rao (1936), Delmalsy (1958), and Bonnet (1957). Moore (1969) gives a general review of the literature on Azolla and indicates its use in agriculture. A total of 85 collections were examined, 70 in detail. When available, at least 10 plants per collection were examined under a binocular microscope. Wetting was sometimes used to bring out detail. Massulae were placed in water under a vacuum for two hours to remove air bubbles. Following this, they were mounted in polyvinyl aleohol (Elvanol solution) on a cover slip which, when dry, was affixed to a slide with Canada balsam. The megaspore complexes were treated with acetolysis solution for 5-10 minutes at 200°F, F, washed, and then oxidized with Shultze solution eins . P. Wolters with the drafting . The manuscript was typed by Miss N. Peis aser. Expenses e covered 7 a National Research Council ‘Grant-in- ‘Aid of Research to L. V. H. Volume 70, No. 4, of the JourNat, pp. 129-172, was issued Jan. 6, 1971. Z AMERICAN FERN JOURNAL + KCIO;). Mounting followed the same procedure as with the massulae. TERMINOLOGY The definition of new terms and terms which have been used as applied to Azolla in conflicting ways are as follows: Trichomes.—The spine-like projections arising from the angles between the contact areas on the proximal surface of the massulae, having vacuoles continuous with those of the massulae (Shen, 1961). The restriction of trichomes to the proximal surface and the distribution of vacuoles differentiates trichomes from glochidia which occur in the species of section A zolla. Megaspore complex.—The composite structure from which the female gametophyte develops. Cap.—The remnant of the indusium persisting as a total or partial covering over the floats (Figs. 1, 2). Supraspore (new term).—The apical region of the megaspore complex (Fig. 1) consisting of: The columella or apical extension of the perine forming a central conical projection (Fig. 6); the columnar filaments or fibrils arising from the apex of the columella and spreading over the surface of the"upper floats; and the floats (Figs. 1, 2). Collar. —The region intermediate between the supraspore and the infraspore formed by a thickening of the perine. Infraspore (new term).—The distal area of the megaspore com- plex (Fig. 1) consisting of the exine or wall of the megaspore or germinal body (Figs. 3, 5, 20); and the perine or wall layer formed exterior to and free from the exine (Figs. 1, Inperine (new term).—The inner, homogeneous or vacuolate zone of the perine located adjacent to the exine (Figs. 5, 20). Experine (new term).—The outer, sculptured zone of the perine. Experine 1 (new term).—The inner zone of the experine with a vacuolate or pillar-like character which acts as a foot layer for the outer elements of sculpturing (Figs. 5, 20). Experine 2 (new term).—The outer zone of the experine com=- posed of the larger elements of sculpturing (Figs. 5, 20). AZOLLA PINNATA 3 The subdivision given for the infraspore agrees in principle with that of Strasburger (1873), Rao (1936), Delmalsy (1958), Bonnet (1957), Kempf (1969) and Snead (1969), but differs from that of Hall (1968), who considers only the outer fibrous layer found on some fossil forms of Azolla to be perisporial material. He includes our experine and inperine as part of the megaspore exine. AZOLLA PINNATA R. Brown, Prodr. Fl. Nov. Holl. 167. 1810. Plants small, floating a as with bilobed leaves. Pubescence oes commonly onto margin of ventral leaf age Ventral leaf lobes (Fig. 15) composed of single layer of cells, 1.5-2 times larger than dorsal leaf lobes. Two or more leaves before os cn branch, erally 9 ae symmetry Bie in groups of 3. ; each . group containing 1 diamond-shaped upper float 1.5-2 times the size of the 2 trapezoidal lower ee (Fig. 21). Sometimes one extra small lower float (Figs. 1, 2). Upper floats covered by columnar filaments. Inperine and experine apparent in columella (Fig. 6). Expansion of inperine at top of infraspore forming indistinct collar (Fig. 6). Infraspore 225-450 u (avg. 3102) wide, composing 14-14 of megaspore complex. Trilete rays 14 length of megaspore Sout exine perforate, 7 w thick (Fig. 3). Perine 16 » thick with eneous inperine 6 u thick (Figs 5, 20); experine 1 of ir ees pillar-like structures, 1-2 thick (Fig. 20); and experine 2 of irregular en an and fusing baculae 2-5 » in meanen 9p rm in viet “— from inperine to which are fuse Swollen baculae (Figs. 4, 20). ecco density variable (Figs. 7-9). Surface “soon rare or tee Microsporangia contai 3-8 m ae each having a convex ing distal and a pyram ee proximal Psrain (Fig. 13). cre 135- 210 u (avg. 190») long, 150 » wide, ane te ee Ep 2-10, usually 4—5 in contr eee 45-150 pu t base; varying in shape from threadlike to ia (Pig. 1 z. 13). ‘Spine structurally resembling short trichomes, up to 204 t the entire ae = (Fig. 12). a se Lia granular or AMERICAN FERN JOURNAL 1 VoutumME 71, Pate 1 prolongation _-experine 2 experine 1 Detaits or MegasporE CoMPLEXES AZOLLA PINNATA 5 PROBABLE ISOSYNTYPES: Iter Australiense, 1802-05, R. dan 134, 135 (Presented by direction of J. J. Bennett, 1876) (E! Rance: As in Plate 4, see also p. 7). The size of the megaspore. complex and component structures was found to be relatively constant within individual collections, but the total inter-collection variability was large. In particular, megaspores complexes from Africa were notably larger (avg. 600 ») than those from other regions (avg. 475). As this size difference was not correlated with any distinctive structural fea- tures either within the megaspore complex or other plant parts, we believe it best not to use it to establish a distinct taxonomic group. Inter-collection variation in the density of the prolongations occurred randomly with respect to geographic distribution; there- fore, it is not taxonomically useful. In 5-10% of the specimens one additional float appears either by division of one of the lower floats (Figs. 2, 22), or as an extra small float at the base of the supraspore either between two floats (Fig. 23), or below and partially replacing one float (Figs. 1, 24). The occurrence of floats supplementary to the basic number of nine may be used in attempts to work out the phylogeny of Azolla (see Hills and Gopal, 1967). One double megaspore complex was observed (Fig. 10) where two infraspores were fused together, the supraspores being absent (see also Strasburger, 1893, Fig. 100). The greatest development of spines on massulae was seen on specimens from Africa. Specimens from Australia and New Guinea had spines only at the corners of lenticular-shaped massulae. Al- though not all massulae from Africa, Australia and New Guinea had spines, this was the only morphologic variation which cor- related in part with specific plant types. In the future this feature Fig. 1.8 SU : MENTARY LOWER FLOAT. FIG. 2, ‘ice i DIVIDED LOWER FLOAT. fis. 3. Mega- SPORE WITH PERFORATE EXINE AND TRILETE MARK. Fic. 4. ENLARGEMENT OF, CONVOLUTED PROLONGATIONS. Fig. 5. WALL CROSS-SECTION. Fie, 6. LONGI-: TUDINAL SECTION OF MEGASPORE COMPLEX. Figs. 7-9. ENLARGEMENTS OF INFRASPORES ILLUSTRATING VARIATION IN PROLONGATION DENSITY. Fie. 10. FUSED INFRASPORES. AMERICAN FERN JOURNAL VOLUME 71, PLATE 2 DeETaILs oF MassuLaE AND LEAVES AZOLLA PINNATA 7 should be noted, as it may be diagnostic. Branched trichomes (Fig. 14) were observed by Rao (1936). The abundance of branching is variable, occurring in about 10% of the trichomes. Within the same collection or even in massulae from the same microsporangia some massulae have branched trichomes, whereas others lack them; hence the presence or absence of branched trichomes is not a reliable taxonomic character. Morphologic variability in the fruiting structures cannot be used to subdivide A. pinnata into varieties. However, on the basis of leaf-branch morphology two varieties can be recognized. AZOLLA PINNATA R. Brown var. PINNATA Azolla africana Desv. Ann. Soc. Linn. Paris 6: 270. 1827. Azolla guineensis Schum. Beskrivelse af Guineiske planter. 462. 1827 [a preprint]. See J gas Dansk Vid. Selsk. Skr. Natur. Mat. Afhandl. 4: 236. 182 Azolla eae var. Beek (Desv.) Baker, Handb. Fern Allies. 138. 188 Plants 1.5-2.5 em long, 1-1.5 em wide, with dominant main axis and age appearance (Fig. 18). Dorsal leaf lobes 0.8-1.9 mm long, 0.5-1.10 mm wide, length-width ratio 1.4-2.1, imbricate to hi hie imbricate (ma be non-imbricate), relatively thick, curled both longitudinally and transversely, elongate, acute at the apex, with a hyaline margin of irregular width formed by up to four rows of elongate cells, the tips protruding to give a serrated hinat pubescence on upper part, this sometimes accompanied by longi- ean sho Leaves "Li6 (avg. 8) before the first branch. : Australia, New Guinea, Africa and Madagascar. Col- ‘action 30 (Pl. 4) from Thailand is most similar to plants from N d may represent a population introduced for agri- cultural purposes; otherwise it is disjunct in its occurrence). COLLECTIONS SEEN: See page 11 and Plate 4. AZOLLA PINNATA var. IMBRICATA (Roxb.) Bonap. Notes Pterid. 7: 130. 1918. Fig. 11. EquaToRIAL VIEW OF MASSULAE SHOWING END SPINES, TRICHOMES E 16. PLANT OF VAR. IMBRICATA. Fia. 17. DoRSAL LEAF LOBES OF VAR. IMBRICATA. Fig. 18. PLANT oF VAR. PINNATA. Fig. 19. DORSAL LEAF LOBES OF VAR. PINNATA. AMERICAN FERN JOURNAL VoLuME 71, PLATE 3 20 SS | ices 5 TT T exin 22 23 24 26 Dane 40 - 40 30 Ss o 8 > a . s z > 20 = w = = MH : 3 - > a sy ie : o 0 Var. pinnata Share | fos eg SON Ve Bs 1.0 14 1.8 22 NUMBER OF LEAVES LENGTH=WIDTH RATIO MorpuHo.oey AND DistTRIBUTION OF A. PINNATA. Fig. 20. ScHEMATIC DIAGRAM OF MEGASPORE COMPLEX WALL MORPHOLOGY. Fig. 21. RBLATIVE SIZE AND SHAPE OF FLOATS. Fig. 22. Lert LOWER FLOAT DIVIDED. Fig. 23. ADDITIONAL FLOAT BETWEEN PAIR OF LOWER FLOATS. Fia. 24. ADDITIONAL FLOAT BELOW ONE LOWER FLOAT. Fig. 25. SCHEMATIC REPRE- SENTATION OF LEAF-BRANCH PATTERN. Fig, 26. DistRIBUTION OF THE NUMBER OF LEAVES BEFORE THE FIRST LATERAL BRANCH. Fiq. 27. DISTRIBUTION OF THE LEAF LENGTH-WIDTH RATIOS. AZOLLA PINNATA 9 Salvinia imbricata Roxb. ex. Griff. Calcutta J. Nat. Hist. 4: 469. 1844. Azolla imbricata (Roxb.) Nakai, Bot. Mag. Tokyo 39: 185. 1925. Plants 1-1.5 cm diameter. Main stem not dominant, but with aleiornae whe type of branching (Fig. 16). Dorsal leaf ‘ohieh 0.8-1.5 mm long, 0.6-1.0 mm wide, length-width ratio 1.1-1.5, enerally non-imbricate to slightly | imbricate, thin, flat, ovate to brapéacidal, rounded to subrounded at the apex, with a hyaline margin of regular width formed by 2-5 rows of nearly equidi- mensional cells (Fig. 17). Echinate pubescence on basal portion of sie leaf lobe grading to evenly distributed conical pubescence upper bet (Fig. 17). Leaves 2-8 (avg. 4-5) before the first laters branch Paina India, Nopal Ceylon, Burma, Southeast Asia, Philip- China, and Japan; one disjunct collection from Africa. Plants intermediate ose var. ptnnata and var. imbricata are found in Australia, New Guinea, Africa and Indi COLLECTIONS SEEN: See page 11 and Plate 4. Azolla pinnata var. imbricata is distinguished by the plants being generally smaller and more circular in shape, with rounded to subrounded, ovate to trapezoidal dorsal leaf lobes, these with a regular hyaline margin of nearly equidimensional cells and regularly distributed conical pubescence. The trivial name imbricata was an unfortunate choice by Roxburgh (1844), who compared them with the non-imbricate Salvinia. To him the Azolla would seem imbricate, as the ventral leaf lobes are overlapping and the dorsal leaf lobes are never widely spaced. However, we have found these plants to be non-imbricate compared to those of var. pinnata. The distinctiveness of the leaf-branch pattern in different species of Azolla was first suggested by Strasburger (1873). Bonnet (1957) states that in A. pinnata 3-9 leaves precede the first branch and the number of leaves before the first lateral branch increases toward the apex of the plant. We observed 2-16 leaves and no consistent increase or decrease towards the apex. Figure 26 shows the distribution of average leaf numbers before the first branch, based on at least 10 counts per collection. Although in general VouuME 71, PLATE 4 AMERICAN FERN JOURNAL 3901 4¥7) vS¥00 B38WNN v9) 2zindv A eS 19 Jv D159! a3aNnnou-ens : ATHOWH 43H10 A tot a $9 -| Jiv>1waWi B 1WDINOD '2IVONOT3 A : fe : F Juve TWDINOD ‘avINOW A ri-ti db 6-8 a e:z + ANHONIS -NON (22v45NS Twdidw B3ddn) 3DN3DS30NG Ouva M/) HONWaS 1S¥14 990198 NOLLW>INeWI voliv "e"4 CHARACTERS AND GEoGrapuic DISTRIBUTION AZOLLA PINNATA 11 fewer leaves occur before the first branch in var. imbricata than in var. pinnata, leaf-branch pattern is a more useful criterion for separating species of Azolla (Bonnet, 1957) than for separating varieties within A. pinnata. Figure 27 shows the distribution of length-width ratio ted the dorsal leaf lobes, based on at least five mature specimens per collection. This ratio, if considered in conjunction with oa criteria, is useful in distinguishing the two varieties. The leaf-branch pattern, the apical shape, the degree of imbrica- tion, the type of pubescence, and the length-width ratio of the dorsal leaf lobe are symbolically represented with respect to. geo- graphical distribution in Plate 4. In the preceding descriptions and in Plate 4 the term “‘non-imbricate” describes leaves in which there is no overlap; “highly imbricate” when they overlap for at ‘least 2/3 of their length, and “imbricate” when the overlap is intermediate. ‘Regular conical” indicates evenly distributed pu- bescence whose length is less than twice the width and “elongate conical” indicates irregularly distributed conical pubescence ac- - companied by longitudinal striations. The numbers in Plate 4 refer to the following collections: Var. PINNATA: 1, R. Brown 134 (E); 2, R. Brown 135 (E); 4,8. Helms 413 (C); 5, M. Clemens 1947 (OSC); 9, Baron Von Mueller (CAL); 11, R. Schodde 2673 (US); 12, L. Brass 5843 (C); 13, A. Harre 270 (F); 14, J. Brown-Lester in 1890-1891 (CAL, E); 15, H. des Abbayes 2251 (US); 16, H. des Abbayes 2252 (US); 18, O. Hagerup 751 (C); 19, H. Baum 106 (M, E); 20, A. Peter 35683 (UC); 21, A. Peter 44388 (UC); 23, R. Schlechter 12033 (M); 24, H. Soyauz, 24/11/1874 (M); 25, H. M. Richards 8313; 26, R. B. Drummond & A. J. Cookson 6272; 27, J. M. Hildebrandt 3065 (M); 28, G. F. 8S. Elliot 1798 (E); 30, Th. Sorensen, K. Larsen & B. Hansen 547 (C). Examined but not plotted: N. H. W. Brown (E), Gower Collection, River Quenso (E). Var. mpricata: 17, P. Cadjoe 983 (M); 32, Em. Bodinier 1427 (E); 33, M. S. Clemens, 3/1907 (M); 34, A. Loher 1149 (M, US); 35, H. M. Curran 11624 (C); 36, A. M. Evans 540 (TENN); 37, G. Murata 27129 (US); 38, © ease in 1842 (M); 39, T. Nakano in 6/1896; 40, M. Hutch 17883 (E); 1, L. H. Bailey in 7/27/1917 (US); 42, A. Henry 3977 (CAL); 43, H. C. Cheo th (E); 44, Kweichow Expedition 4220 (C); 45, H. Smith 1518 (C); 46, Th. ensen, K. Larsen & B. Hansen 1829 (C); 47, Urupai on 2/21/1969; 48, J. Sinclair on 12/12/1948 (E); 49, J. Sinclair on 14/11/1950 (E); 50, Fer- guson (M); 51, Thwaites C. P. [Ceylon Plants] 1421 (B);. 52, Marshall Ward 12 AMERICAN FERN JOURNAL in 10/1880 (E); 53, A. O. Brodie in 9/186? (E); 55, C. E. C. Fischer 4027 (CAL); 56, Dr. Wight in 1831 (E); 57, Shack Mo Kim in 1896 (CAL); 58, P. M. Debbarman 644 (CAL); 59, S..Kurz 8194 (CAL); 60, J.D. H. (C); 61, J.J. Wood 187 (CAL); 62, C.. Maries 1890 (CAL); 63, H.H. Johnston on 1/27/1896 (E); 64, O. Polunin, W. R. Sykes & L. H. J. Williams, 1822 (E); 65, Stainton, Sykes & Williams 6952 (E); 66, D. H. Nicolson 2911 (US); 67, Stainton, Sykes & Williams 2546 (CAL, E); 68, East India, Dr. Wight (E); 69, H. Collett, 11/1890 (CAL); 70, @. Watt 5090 (CAL). Examined but not plotted: M. Vanoverbergh 3226 (BKL); R. R. Stewart 1359 (BKL); Gower Collection (E); Wight 5 (C). INTERMEDIATE TyPEs: 3, J. Camfield in 4/1897 (US); 6, N. Michael 1431 (E); 7, W. Layer (M); 8, R. Specht 1166 (US); 10, P. Darbyshire 803 (US); 22, H. Merzmuller & W. Guiess on 8/03/1958 (M); 29, J. M. Hildebrandt 35965 (M); 31, G. H. Vachell on 17/7/1830 (E); 54, Stocks, Law (C). The distribution of the two varieties presents an interesting problem. The close similarity of the disjunct populations of var. ptnnata indicates the populations were once connected. Mega- Spore complexes similar to those of A. pinnata in Upper Pliocene- Lower Pleistocene strata of Russia (Nikitin, 1957 ; Dorofeve, 1963) indicate the species was once much more widespread. It is therefore reasonable to assume that the separation of popu- lations of var. pinnata may have occurred as the result of glaciation during the Pleistocene (ca. last 3 million years). Plants typical of var. pinnata were not seen in collections from India and Ceylon. However, some plants from these regions had notably higher leaf numbers before the first branch than usual is for var. imbricata. Other plants had pubescence more like that of var. pinnata. This may suggest a former relic population of var. pinnata in India and/or Ceylon which has influenced the character of, and has been absorbed into, the present population. Attempts were made to relate the distribution of the two vari- eties to generalized plots of variations in temperature and light intensity. No direct relationship between these factors and the distribution pattern was found. It was noted that var. imbricata occurs in more densely populated areas, suggesting that some agricultural practice may influence the distribution. However, the succession of plants from Australia to Southeast Asia appears to reflect a clinal development, and therefore agriculture is probably not the controlling factor. AZOLLA PINNATA 13 Future paleobotanic studies will unquestionably add to our knowledge of the former distribution of the species. The answer to the relationship between the two varieties and the reasons for the distinct geographic ranges will best be answered by the study of the varieties themselves. Future collections should be more specific as to the character of the collecting site and its ecological setting. LITERATURE CITED Baker, J. G. 1887. Handbook of the fern allies. George Bell, London. Bonnet, A. L. M. 1957. Contribution a l’étude des Hydoptéridées, III. pom ne sur Azolla filiculoides Lamk. Rev. Cytol. Biol. Végét. -88. Ceeiadcs C. 1905-6. Index Filicum. Hagerup, Copenhag Detma.sy, P, 1958. Nouvelles recherches sur le eat d’Azolla. La Cellule 59: 235-268. Dororery, P. I. 1963. The Tertiary — of Western Siberia, Akad. Nauk SSR, Bot. Inst. Komarov, Len Grirritu, W. 1845. On Azolla and Salvinia, Clee T. Nat. Hist. 5: 227-273, Hau, J. W. and N. P. Swanson 1968. Studies of fossil Azolla: Azolla mon- tana, a Cretaceous megaspore with many small floats. Amer. J. Bot. 55: 1055-1061. Hints, L. V. and B. Gora. 1967, Azolla primaeva and its phylogenetic parm Canad. NB Bot. 45: 1179-1] 191. Kemper, E. K. 1 1 orodermis von kanozoischen Megasporen der Wasserfarn-Gattung rey Paliont. Zeitschr. 43: Moorg, "e oe 1969. Azolla: Biology and agronomic significance. Bot. Rev. Nixitin, P. A. 1957. Pliocene 08 Quaternary floras from Voronezh District. Acad. Nauk SSSR, Bot. Inst., Moscow—Leningrad. Rao, H. 8. 1936. The structure re life history of liao pinnata. R. Br. with poms on the fossil history of the Hydropterideae. Proc. Indian Acad. Sci. 2: 175-200. Suen, E. Y. F. 1961. Concerning Azolla imbricata. Amer. Fern J. 51: 151-155. SNeap, G. 1969. Microfloral —— of the Cretaceous-Tertiary bound- ry, Central Alberta. Res. Coun. Alberta Bull. 25. Secs E. 1873. Ueber Aa Friedrich von Zezschwitz, Jena. Sup, 8. R. 1934, A preliminary note on the study of Azolla pinnata R. Br. J. Indian Bot. Soc. 13: 189-197. DEPARTMENT oF GEOLOGY, UNIVERSITY OF CALGARY, CALGARY 44, ALBERTA, CANADA. 14 AMERICAN FERN JOURNAL A New Variety of Lindsaea odorata Roxb. from Darjeeling in the Himalayas T. SEN AND U. Sen! During our field studies in Darjeeling in 1965 and 1967, discovered some Lindsaea specimens that were strikingly flank from the species of Lindsaea familiar to us. The plants resemble Lindsaea odorata Roxb.,? which is also found in Darjeeling. How- ever, close examination showed that they are quite distinct: they are smaller and have short branched foliar hairs, but lack multi- cellular long hairs and internal endodermis in the rhizome. These plants thus represent an undescribed taxon. LINDSAEA OporRATA var. darjeelingensis Sen & Sen, var. nov. A var. odorata differt: frondibus multo minoribus, usque ad 15 cm longis et 1.7 cm latis; paleis Siberintis latitudine basi ex cellulis 4-7 constantibus; frondibus juvenilibus pilis brevibus ramosis praeditis; endodermide interiore stelis rhizomatis carente. Rhizome creeping, short, slender, ca 2 mm i Raycom scales deep. brown, acuminate, up to 7 ¢ ells broad at the base; vascular 2-fo 1-4 veins ending at the margin of each lobe; laminae of young fronds with small, 2- or 3-c elled, unbranched, glandular hairs and 3-celled branched glandular hairs; sporangia globose; annulus of 15-25 cells; stomium 8-12 cells; paraphyses present with sporangia ; spores bilateral, 52 u X 38 u X 33 mu, enclosed by a thin perine, this cracking off on aretolyii. Type: Darjeeling, on way to Birch Hill, Sen & Sen 505 (holo- type, Diaries of Botany, University of Kalyani, West Bengal). 1 We wish to thank Professor R. E. Holttum for the Latin diagno s, and Professor S. P. Sen for his caged es ent. See pn aie : * Kr a 15: 567, 1967) has shown this to be the correct name for what has always, but Eeomeiie: been called Pas aie cultrata (Willd.) Swartz. AMERICAN FERN JOURNAL VoLuME 71, Puate 5 LINDSAEA ODORATA var. DARJEELINGENSIS. Fic. 1. ADULT PLANT, X 1. Fig. 2. Ruizome SCALE, X 53. Fig. 3. Pinna, X 5. Fic. 4. UNBRANCHED FOLIAR HAIR, X 140, Fig. 5. BRANCHED FOLIAR HAIR, X 140. Fie. 6. SURFACE VIEW OF EPIDERMAL CELLS ON THE ABAXIAL SIDE OF LAMINA, X 87. Fig. 7 TRANSVERSE SECTION OF THE RHIZOME SHOWING THE ABSENCE OF INNER ENDODERMIs, X 300. Fic. 8. Sporn, X 435. 16 AMERICAN FERN JOURNAL Lene SPECIMENS SEEN: Darjeeling-Sudrapong Road, Sen & Sen 506; Rangaroon Forest, Sen & Sen 507; Lebong, Sen & Sen 508 ; jade Nora G. King BSI 5341, 5844 (C AL). This new variety somewhat resembles var. pallens Hook. (Sp. Fil. 1: 204. 1846) from Nepal and Sylhet in frond size, and yet differs in other characters. The features which distinguish and characterize var. pallens are: (i) the scales are less than 4 cells wide at the base, (ii) short branched hairs and multicellular hairs are seemingly absent even on the youngest fronds, (iii) the vas- cular cylinder of the rhizome has both the inner and outer endo- dermis, and (iv) the spores measure 61 u X 44u X 48 p. We con- clude that var. darjeelingensis is a very distinct local variety of this wide ranging species. The differences among the three varieties of Lindsaea odorata may be summed up in the following key: Scales 2-3 or very rarely 4 cells broad at the base; short branched hairs ab- on the laminae; stele of the rhizome with both inner and outer endodermis. Long multicellular hairs always present on young fronds; fronds ca 60 em long and ca 35mm wide at the broadest region; laminae thick; spores 644 X 44u X 43 nu. var. odorata Long multicellular hairs never occur even on the young fronds; fronds ca 20 cm long and 24 mm wide at the broadest region; laminae thin; spores 61 » X 444 X 484 var. pallens Seales 4-7 cells broad at the base; short beached hairs always fase on young laminae; stele of the rhizome with only the outer endodermis; long multicellular hairs never develop on the fronds; fronds ca 15 em long and 17 mm wide at the broadest region; spores 52 X 33 var. darjeelingensis DEPARTMENT OF Botany, Kanyani UNIvERsSITY, Ka1yANnt, West Benaat, Inpia. MENISCIUM MACROPHYLLUM 17 The Proper Disposition of Meniscium macrophyllum Kunze C. V. Morton In our paper “The American Species of Dryopteris, Subgenus Meniscium” (Bull. Torrey Bot. Club 65: 347-376. 1938), Dr. Maxon and I concluded that Meniscitwm macrophyllum Kunze could not properly be referred to Dryopteris subg. Meniscitum because some of the sporangia are borne on the leaf surfaces be- tween the veins, thus being ‘‘acrostichoid.’’ Consequently, we proposed to refer the species to Bolbitis, where the acrostichoid condition is tne rule. However, I have never been really satisfi with this disposition, because this species resembles species of the subg. Menisctum much more than any Bolbitis species in most characters. My reexamination of the available specimens indicates that ‘in some specimens the sporangia are borne only on the arcuate cross-veinlets and also, exceptionally in Meniscium, on the short, medial, outwardly excurrent veinlets also. Indeed it appears that in semifertile blades the sporangia are mostly on the excurrent veinlets (ef. Spruce 3030). In some specimens the sporangia do extend out also onto the leaf surfaces, which is certainly a unique character in Meniscium but not sufficient to necessitate the reference of this species to a different genus, especially when the character is not shown by all specimens. Therefore, I now refer M. macrophyllum to Thelypteris subg. Meniscium: THELYPTERIS macrophylla (Kunze) Morton, comb. n Menisciu'm macrophyllum Kunze, Flora 22(1): Beibl. 44. ak: “Type: Serra o Mar, Bahia, Brazil, Martius 363 (isotypes BR, K, NY). PAlccdies meniicieldes Fée, Mém. Foug. 2: 93, ¢. 55. 1845. Tyre: Bahia, Brazil, Blanchet (not seen). I have seen two specimens ¢ collected by Blanchet in Bahia, numbers 2228 and 2477, both at Kew, and these are very likely authentic and parts of the type collections ?Poecilopteris meniscioides (Fée) K. B. Presl, Epim. Bot. 175. 1849 [1851]. Phegopteris macrophylla (Kunze) Mett. Fil. Lechl. 2: 24. 1859, at least as to basionym. Nephrodium macrophyllim (Kunze) Keys. Pol. Cyath. Herb. Bung. 48. 1873. Dryopteris macrophylla (Kunze) C. Chr. Ind. Fil. Suppl. 1: 35. 1913. 18 AMERICAN FERN JOURNAL Bolbitis macrophylla (Kunze) Maxon & Morton, Bull. Torrey Bot. Club 65: 375. 193 ApDITIONAL SPECIMENS EXAMINED: Braziu: San Isidro, Goyaz, Pohl (BR). Limocero, Goyaz, Pohl (BR). Serra do Mar, Bahia, Luschnath' (BR). Theos, Babia, Luschnath! (BR). Without locality, Riedel (BR, sterile). ee AME: Jodensavanne-Mapanne Creek, Suriname River, Lindeman 45 (U); Hekking 1212 (U). Blanche Marievallen,; Nickerie River, Maas & eae 10987 Britisa GUIANA: Aipien 782 (K). In our discussion of this species, Maxon and I decided that specimens with setose sporangia ought to be referred also to B. macroyylla, but I have now changed my mind about that also. The type of M. macrophyllum and other Brazilian specimens that I have recently examined (but not all of those cited in 1938 have been reexamined) have the sporangia non-setose. Those from Trinidad and Tobago and some of those from the Guianas have them conspicuously setose, and in a peculiar way. The setae, numbering from one to four, are exceedingly conspicuous and are borne on the annulus. In the other species of Menisctum with setose sporangia we stated that the setae are borne on the stalk of the sporangium. I am now unable to verify this in all cases. Sometimes the seta 7s on the sporangium stalk, but at other times it seems to be only from the receptacle rather than the sporangia, thus being technically a paraphysis. It may be that these setae are too easily detached and are separated from their sporangium readily in making a microscope slide. However this may be, it does seem that the character is still a usable one in separating those species with setose sporangia or paraphysate sori from those with glabrous sporangia and sori. Since I have used this char- acter of setose sporangia as a fundamental character in this group, it is not reasonable to allow both types within T. macrophylla, especially since there is some geographic disjunction and some other characters of more debatable importance. It appears that in typical T. macrophylla the excurrent veinlet is well developed ‘ These collections may very well be isotypes, for the specimens distributed Martius in his numbered sets were often collected by others, among them ? uschnath, They are in any case topotypes and quite ‘deskion! to Martius 363. MENISCIUM MACROPHYLLUM 19 in the fertile pinnae, and it bears sporangia. In the plants with setose sporangia the arcuate cross veinlets are nearly straight and the excurrent veinlet is generally reduced to a mere knob hardly long enough to bear sporangia. This character ought to be checked in the field. I do not have enough good fertile material. This plant with setose sporangia may be known as: THELYPTERIS a bapisniog (Fée) monies cont, no Meniscium guyanense Fée, Gen. Fil. 224 2. TYPE: ee in sylvis paludosis, ad amnes Conana et cette ” French Guiana, Leprieur in 1835 (originally determined as M. oorbifolecn’ (P?, not seen). The description, especially the statement that the sporangia are ilose, indicates that the name refers to this species. ale sna fendleri Baker, Journ. Bot. Brit. & For. 25: 100. 1887. Type: nidad, Fendler 88 (presumably K, not seen; isotypes NY, US). Wikies tibitiets Hort. ex Baker, eS Bot. 5: 487. 1891. Lectotype: Hort. Linden, ex Herb. Moore (K). A specimen “‘e caldario Linden”’ in Brussels is probably an isolectotype (Weatherby photograph US). A later collection, one from authentic material, is cult. Jan. 7, 1871, ex Herb. R. Sims (K). Probably this name ought to be con- sidered as invalid, since it was not definitely accepted as a valid ecies by the publishing author, Baker. Leptochilus fendleri (Baker) C. Chr. Ind. Fil. 385. 1906. Based on Acrostichum endlert Baker (not M a fendleri Eaton?). Diyos anceps Maxon, Contr. U.S. Nat. Herb. 24: 62. 1922. Based on Acrostichum fendleri a not Dryopteris fendleri (D. C. Eaton) Kuntze (1891 I have not been able to reexamine all the specimens cited in 1938. Those from the Guianas should be rechecked, since both 7. macrophylla and T. guyanensis grow there. SPECIMENS oF T. GUYANENSIS EXAMINED: * In the “Index Filicum”’ the citation for this combination is given as Chr. Bot. Tidsskr. 26(2): 285. 1904, but at this place it is given as merely Peploctilies fendler ri (Eat.), with no basionym cited. There is, t e that could be a basionym. As shown in “Inde wae nae er “ihe e impression that the name Meniscium fendleri MS and that Acrostichum fendleri Baker was a Bide er ah tage Senin e cit aton never published such a name at the p ae a Aa oa: 62. 1922, a new spe ow ens nder feseay hilus fendleri i in the ‘Index ith the change of parenthetical author find Eaton to Ba 20 AMERICAN FERN JOURNAL TRINIDAD: Orange Road, Hombersley 226 (US). Brazil, Britton, Briiton & re 2121 (US); Withou: locality, Fendler 88 (NY, US); Bot. Gard. Herb. 584 (US). oak AGO: Between Roxborough and Parlatuvier, Broadway 4904 (US) Surinam: Julianatop, Wilhelmina Mountains, 500 m., i es 10451 (U). Maratakka-Saparra, Gonggrijp & Stahel 971 (U, US). Bakhuis Mountains, between Kabalebo River and Left Coppename River, Pasechaite & Maas 3054 (U). Britisn Guiana: Barina River, Jenman (NY, 8 sheets). Demerara River, Jenman (NY). These two species, 7’. macrophylla and T. guyanensis, are most likely to be confused with 7. chrysodioides (Fée) Morton and its variety goyazensis, which also have the fertile leaves smaller than the sterile, the blades being therefore subdimorphic. The other large species that might be confused are included in the following key: Sporangia non-setose. Sporangia confined to the arcuate cross-veinlets; bases of pinnae broad, t cuneate. Pinnae broadest at base (Florida, West Indies, Margarita Island yehenaeln i oe T. reticulata (L.) Proctor Pinnae broadest above base (Costa ee T. turrialbae (Rosenst.) Morton saeieaap tt on the arcuate veinlets, the a veinlets, and some- es also on leaf tissue; pinnae cuneate at base (Brazil, Peru, Golteas Amazonlan Venezue la). T. macrophylla (Kunze) Morton Sporangia setose on the stalk or annulus Sporangia setose on the annulus;? arcuate tanga potas straight, the excurrent veinlet often reduced to a mere knob (Trinidad, Tobago, Guianas)............ gum mensis (Fée) Morton ra setose only on the sporangium stalk excurrent veinlet well developed, not bearing sporangia (Guianas, Braz chrysodioides (Fée) Morton var. chrysodioides! T. ¢. var, goyazensis (Maxon & Morton) Morton Nationa! "Museum or NaTurAL History, WAsHinaTon, D. C. I : the sae are ea is sometimes not easily seg rer gti It is likely that mature material. chrysodioides is accepted here in our 1938 hi but wh Sith correctly or not is not Salts ometuae until Fée’s THELYPTERIS NORMALIS COMPLEX 21 The Thelypteris normalis Complex in the Southeastern United States! A. Rep SMITH The species of the Thelypteris normalis complex in the south- eastern United States and the West Indies have long been acknow- ledged as taxonomically troublesome. The first person to mono- graph the group was Christensen (1913), who treated it under Dryopteris, subg. Cyclosorus. More recent workers have also recognized the integrity of the group, but generally have treated it as a subgroup of Thelypteris (Small, 1938; Wherry, 1943, 1964). The chief problems encountered -by Christensen and others have been the extreme variability in the species and the large number of “forms” which are seemingly intermediate between species. Robert and Edward §t. John, avid collectors of Florida ferns in the 1930’s, added to the- proliferation of names when they described numerous Florida ‘“endemics.’”’ The assiduousness of previous workers, however, has laid the groundwork for more detailed studies utilizing cytology and field-to-greenhouse trans- plants, an approach which I believe has clarified the systematics of the group. A revision of Thelypteris sect. Cyclosorus in the New World is currently being prepared for future publication. However, in order that the important conclusions of this study might be more immediately available to those with an interest in ferns of the southeastern United States and to those involved in the prepara- tion of floras, it seems appropriate to publish a portion of the study dealing with the species of this area. Following a key to the species and varieties is a synopsis of the southeastern United States taxa, excluding 7. totta (= T. gongylodes), which is only rather distantly related to species in the 7’. normalis complex. Important aspects of synonymy, habitat, distribution, and a discussion of 1 Much of this research was conducted under a National Defense Education Act Fellowship. a2 AMERICAN FERN JOURNAL the salient differences among the taxa are given, together with a table summarizing the cytological findings of this study. Docu- mentation of chromosome numbers will be presented elsewhere. Key TO THE SPECIES AND VARIETIES OF THELYPTERIS SECT. CyYCLOSORUS IN THE SOUTHEASTERN UNITED STATES 1. Basal veins of Sa at segments united below the sinus with an excurrent ward the sinus; costules, veins, and often lamina above ha 2. Rhizomes piie-eheoping: coe nearly naked; costae below with light- colored scales; lamina, veins, costules, aad costae above glabrous, or sparsely Dubesoent with very short hairs. 3. Lamina and veins pubescent below; indusia and costal scales h t SV So i Po. Pe ie on T. lotta var. hirsuta 3. Lamina and veins glabrous — indusia and costal scales glabrous or with sparse hairs............ T’.. totta var. totta 2. Rhizomes pen or pieced costae’ below without scales; adaxial surface of lamina glabrous or hairy; veins, costules and costae above moderately to rather densely hairy with rather long stout hairs. 4. Costae below with predominantly short hairs uniform in length (less than 0.2 mm and usually less than 0.1 mm long); excurrent veins mostly greater than 2 mm long; stipes purplish; feiide with usually more than two ee of sires reduced pinnae at the base................ E, entala 4. Costae below with most hairs greater than 0.3 mm long gears ex- c g 0.5 mm), not uniform; excurrent veins less than 2 mm long; stipes stramineous; eee: with 0-2 pair(s) of slightly reduced pinnae at the bas he cattnen a var. versicolor 1. Basal veins of adjacent segments free below, or connivent at the sinus; costules, veins, and lamina above with or without hairs 5. Rhiscnes: suberect to erect, massive; basal segments of the lower and ial pinnae usually narrowly elongate parallel to the rachis; pa (enlarged basal acroscopic pinna segments) often pre- oe = the base of the lower pinnae; costae below without Se See fe OO ak ANON Cae ier patens var. patens 5. Vhiscenie long-creeping to short-creeping, hie stipes arcuate at the base; basal segments of the lower and medial pinnae either reduced, the same size as, or slightly longer than more distal pinnules; auricles small or absent; costae below with or without small scales. THELYPTERIS NORMALIS COMPLEX ps 6. Costae, een and veins above with at least a few rather stout mostly greater than 0.3 mm long; lamina above es rather glandular; scales absent on mature fronds on rachis and costae below 7. One or two pairs of basal pinnae somewhat reduced; rhizomes short-creeping, sometimes appearing suberect; vena- tion variable, even on the same frond, from anas mosing with a short excurrent vein to connivent at the sinus; lamina above often somewhat hairy; veins above always ies stout hairs, many greater than .4 mm long.. . quadrangularis var. versicolor a: cnn pinnae eau ‘hot reduced; rhizomes short- eeping to long-creeping; veins connivent at the sinus or the distal one of each pair meeting the margin slightly above the sinus; lamina above glabrous or sparsely ee veins above with. or a agar long SOUL HAI rs ee er ee 4. ormalis 6. Costae, costules, oe veins above glabrous or ae very thin short hairs mostly less than 0.2 mm long; lamina above ‘ine a few scales persistent on rachis and sometimes costae below one in T. ovata var. ovata). 8. Terminal ‘‘pinna”’ usually at least 5 times as long as at scales on costae below often numerous; lamin coriaceous; lower pinnae (at their widest point) spa less than 1.2 em wide, incised 0.5-0.7 of their width. 5. T. augescens 8. ud cue ‘“‘pinna’’ less than 5 times as long as wide; scales ostae below absent or relatively sparse; lamina Pieeenerns to subcoriaceous; lower pinnae (at their widest point) usually greater than 1.2 em wide, incised 0.6 to often more than 0.8 of their width. 9. Scales usually absent on the costae below; ey above PIATOUR er 6. T. ovata 9. A few very narrow scales usually present on the costae ; lamina above minutely pubescent or glabrous 6a. T. ovata var. lindheimerit 1. THELYPTERIS DENTATA (Forssk.) E. St. John, Amer. Fern J. 26: 44. 1936. oscar da dentatum Forssk. Flora Aegypt.-arab. 185. 1775. HoLotyps: tains of southwestern Arabia, Yemen, Forsskél s.n. (C-Herb. ForsskAlei 809! photographs K, UC!) 24 AMERICAN FERN JOURNAL Polypodium molle Jaeq. Coll. 3: 188. 1789, non Schreb. 1771; nec All., 85. Hotoryre: cultivated in the Botanical Garden, Schoenbrunn, and said (probably incorrectly) to originate from Caracas, Venezuela (W! photograph UC!). Aspidium molle Swartz, J. Bot. Schrad. 1800(2): 34. 1802. To be con- sidered a new name under Art. 72 (Note) for Polypodium molle Jacq., 1789. Aspidium violascens Link, Hort. Berol. 2: 115. 1833. Hotoryps: cultivated in the Botanical Garden, Berlin, from plants said (probably incorrectly) to originate from Brazil (B, not seen; isotypes HBG! photographs UC! US!; L! photograph UC! US)). Deore mollis (Swartz) Hieron, Hedwigia 46: 348. 1907. Dryopteris dentata (Forssk.) C. Chr., Dansk. Vid. Selsk. Skr. VIII, 6: 24. 1920. Thelypteris reducta Small, Ferns Southeastern States. 254. 1938. LectoTyPE (chosen herein): ‘‘Inter Bowling Green et Fort Green [Hardee Co.], Florida”, 7-11 Dec. 1934, E. St. John, R. St. John, W. A. Knight, and J. K. Small, s.n. (NY!). HapitaT AND DistriBuTION: Roadsides, pastures, edges woods, and other disturbed habitats; also limestone sinks. Tropical and subtropical North and South America, with the possible —— of the Andean countries. This species is a common ‘weed” of waste areas in tropical Africa, Asia, and the Pacific Islands. In Florida, T. dentata is probably confined to hammocks, especially those of a limestone nature. There is the strong indication that this species is not native to the New World but has been introduced rather recently (Smith in Strother and Smith, 1971). The earliest United States collection I have seen is from near Mobile, Alabama, in 1904. It was probably not collected in Florida until 1930, when it was found in Her- nando County. Subsequently, it has been found in at least ten Florida counties and appears to be increasing its range, both in Florida and throughout the New World. Single collections I have seen from Georgia, Kentucky, and Louisiana are probably all escapes from cultivation. That T. dentata has escaped from culti- vation is not surprising, since it is a very common fern in green- houses in the United States. Its native relative, T. quadrangularis, is rarely cultivated. THELYPTERIS NORMALIS COMPLEX 25 From T. quadrangularis, T. dentata differs in the longer excurrent vein, the uniformly short pubescence below (long or long and short hairs mixed in 7. quadrangularis), and the purplish color of the stipe and rachis (stramineous in T. quadrangularis). Thelypteris dentata has been counted many times as tetraploid (n = 72) from both Old and New World collections, while quadrangularis is diploid. 2. THELYPTERIS QUADRANGULARIS (Fée) Schelpe var. versicolor R. St. John) A. Reid Smith, comb. nov. Thelypteris macilenta EB. St. John, Amer. Fern J. 26: 50, 52, pl. 5. 1936. Ho.otryre: 7 mi NW of Brooksville, Annuttalagga Hammock, Florida, E. St. John 204 (NY! photograph UC!; isotypes MO!, 2-NY!, US!). Thelypteris versicolor R. St. John in Small, Ferns Southeastern States. 250, cum tab. 1938. Hoxioryrr: Brooksville, Hernando Co., Florida, R. Si. John 109 (NY! photograph UC!). Thelypteris X versicolor R. St. John (pro sp.). Stat. emend. in Wherry, Southern Fern Guide 120. 1964. Hasirat aNp DistrrBuTiIon: Woods and limestone sinks along the southeastern coastal plain from South Carolina to eastern Texas; also known from Cuba. Other varieties of T’. quadrangularis are found in Mexico, Central and South America, Africa, and probably Malesia (Holttum, pers. comm.). Although previously considered either a species or a hybrid between 7’. dentata and T. normalis, T. versicolor is best treated as a variety of the widespread 7’. quadrangularis, differing from the type of 7. quadrangularis (from French Guiana) by its conni- vent or weakly united veins, short-creeping rhizome, and distri- bution. Typical 7. qguadrangularis from the Lesser Antilles, and Central and South America has united basal veins and an erect rhizome. In the West Indies, var. versicolor grades into two other varieties of 7’. quadrangularis, including the type variety, making specific recognition impossible. Wherry (1964, p. 120) treated 7. versicolor as a hybrid, stating that the sori tend to be “sparse and poorly developed, and [the] 26 AMERICAN FERN JOURNAL spores mostly aborted. [Thelypteris versicolor is| Manifestly of hybrid origin between distantly related taxa,’’ meaning 7’. dentata and T. normalis. I have examined spores of versicolor throughout its range and find that they are normal in appearance. Chromosome counts from Louisiana and Florida (Table I) show that var. versicolor is a diploid with 36 pairs of chromosomes at meiosis. This contrasts with the tetraploid condition in both T. normalis and 7’. dentata. Thelypteris macilenta, known only from the type and from specimens propagated from the original clone, appears distinct but is no more than an aberrant element within var. versicolor The rhizome habit, small size of the fronds, reduced pinnae below, pinna shape, and stout hairs on the ultimate veins and costules both above and below all suggest this relationship. 3. THELYPTERIS PATENS (Swartz) Small, Ferns Southeastern States 243. 1938. Polypodium patens Swartz, Prodr. 133. 1788. LecrotyPE (chosen herein): amaica, Swartz s.n. (S-PA! photograph UC!). Aspidium stipulare Willd., Sp. Pl. ed. 4, 5: 239. 1810. Hotoryre: Plumier, Tract. Fil. pl. 23! Aspidium macrourum Kaulf. Flora 6: 365. 1823. HoLoryre: Martinique, Sieber 354 (not seen; isotypes L, photograph US!, M!, MO!). Nephrodium albescens Desv. Mém. Soc. Linn., Paris 6: 258. 1827. HoLoryPe: Jamaica, collector unknown (P, not seen, photograph US!). Hasirat anp Distrisution: Common along roadsides and paths, edges of woods, and open woods in the West Indies, southern Mexico, Central America, and south to southern Brazil and Bolivia. From the United States I have seen only a single specimen: Florida, Dade Co., § of Miami, Feb.-Mar. 1905 A. A. Eaton - s.n., (US!). Thelypteris patens var. patens can be distinguished from most New World taxa in Cyclosorus by its erect rhizome and by the stipe base scales, which are ovate-lanceolate, light brown, and glabrous. The basal pinna Segments in var. patens are either elongate and acuminate, or the acroscopic segment is expanded THELYPTERIS NORMALIS COMPLEX 27 into a deeply pinnatifid auricle up to 5 cm long. Without the rhizome, a specimen of 7’. patens is sometimes difficult to distinguish from specimens of 7’. normalis. The latter species has hairs on the costules and ultimate veins above and the lamina above is fre- quently glandular. The blade above in T. patens var. patens is glabrous except for hairs along the costa. All specimens of 7’. patens thus far counted have been tetraploid (n = 72). 4. THELYPTERIS NORMALIS ee Chr.) Moxley, Bull. So. Calif. Acad. 19: 57. 1920 Dryopteris patens var. glandulosa A. A. Eaton, Bull. Torr. Club 33: 477. 1906. HoLotryrs: Colwell’s Hammock, Dade Co., Florida, A. A. Eaton 185 (GH! photographs MO!, UC! Dryopteris normalis C. Chr. ae for Bot. 9(11): 31. 1910. SynrypeEs: Jamaica, Jenman Herb. s.n. (NY Thelypteris sazatilis R. St. ei in Small, Ferns Southeastern States 236. 1938. Hotoryre: Ellaville, along the Suwanee River, [Madison Co.], Florida, R. St. John 86 (NY!). Thelypteris unca R. St. John in Small, Ferns Southeastern States. 246. Hoxoryre: Costello Hammock, Dade Co., Florida, R. St. John 172, 2 sheets (NY! photograph UC)). Thelypteris macrorhizoma E. St. John, Amer. Fern J. 32: 146. 1943. Hoo- TYPE: Sheep Island, The Cove, 6 mi from Floral City, Citrus Co., Florida, E. St. John 1273 (FLAS! photograph UC!; isotypes GH?, US! photograph UC)). Hasitat AND DistripuTion: Common in woodlands, limestone sinks, gullies, and wet roadsides in open or partially shaded situ- eastern Texas. Bermuda, Bahama Islands, Greater Antilles, Tortola, and from Costa Rica north through the Yucatan peninsula to southern Tamaulipas, Mexico. Rare in the Lesser Antilles, northern Venezuela, and northeastern Brazil. Reports by Morton (1967) that it occurs south to Bolivia are dubious. Thelypteris normalis is a -well-characterized species but it has been either too broadly or too narrowly construed by most taxonomists. It can best be distinguished from 7’. ovata by the usually hairy costules and veins above, and by the stipitate glands 28 AMERICAN FERN JOURNAL on the adaxial surface of the lamina. In general, the pinnae of T. normalis are not so deeply incised as those of T'. ovata and the segments are less oblique. Sori in 7. normalis are medial to supra- medial, whereas in 7’. ovata they are supramedial to submarginal. The presence of single-celled glands on the sporangial stalks and the absence or paucity of glands on the indusial margin are excel- lent characters for distinguishing 7. normalis from T’. ovata, where the situation is reversed (glands always absent on sporangial stalks but present and often abundant on the indusial margin). Throughout most of its range, 7. normalis is easily distinguished from T. quadrangularis by venation, but in the southeastern United States, T. guadrangularis var. versicolor often has the basal pair of veins only weakly united or merely connivent at the sinus. Thelypteris normalis can then best be identified by the lowermost pinnae being usually not reduced (the lamina deltoid rather than ovate-lanceolate in outline), by the lack of hairs on the leaf tissue between the veins above, by its generally longer creeping rhizome, and by the slightly thicker blade texture. Thelypteris normalis hybridizes with T. augescens, T. ovata var. ovata, and probably T. quadrangularis, although hybridization with the last-named taxon has not yet been cytologically verified. In all cases examined the resulting hybrids are sterile. Morton (1967) attempted to show that there are at least two names with priority over Dryopteris normalis C. Chr. One of these, Aspidium germanii Fée, I have placed in synonymy under Thelypteris quadrangularis. Unfortunately, the type of the second, Nephrodium kunthii Desv., has not been available for study, but I believe that it very likely is not 7. normalis, but rather a variety of T. patens. Thelypteris saxatilis and T. unca are both easily included within the latitude of variation of T. normalis. Thely;teris saxatilis is a somewhat smaller form than is usual for the species; some of the Specimens so labeled by St. John are referable instead to 7’. ovata rather than T. normalis. Thelypteris wnca differs from most speci- mens of T. normalis mainly in its slightly more faleate pinnules. Thelypteris macrorhizoma is less pubescent than ‘typical’ normals, THELYPTERIS NORMALIS COMPLEX 29 but differs in no other significant way. 5. THELYPTERIS Soerprrets Dorn Munz & Johnston, Amer. Fern, 122:75-1 Aspidium augescens Link, Fil. Sp. 103. 1841. Type. Cuba, Otto 89, right hand plant (B! photograph UC!; additional authentic material is at BR! and UC!). Cultivated in the Botanical Garden, Berlin, from plants said to. be from Caracas. Christensen (1913) discussed the origin of the type material and concluded, no doubt correctly, that the type locality must have been Cuba. HapitaT anp DistrisutTion: Dade Co., Florida, Cuba, and on the islands of Andros and New Providence in the Bahamas. Generally on exposed limestone outcrops in moist to moderately dry open sites. Thelypteris augescens can be distinguished readily from its close relative T. ovata by the more numerous paleae on the costae below, by the more coriaceous blade, by the greater pinna length/ width ratio, by the less deeply incised pinnae, and by the usually distinct terminal “pinna.’’ Glands are absent on the sporangial stalks, but usually present on the indusial margin, as in 7. ovata. The West Indian 7. serra greatly resembles T. augescens but can be distinguished by its even more abundant costal scales and more distinct terminal “pinna.” Additionally, the basal segments of the lower pinnae in 7’. augescens are never reduced as is usually the case in 7’. serra, but are as long as or slightly longer than the more distal pinnules. Reports of 7. serra in Florida (Small, 1938) are the result of confusion with 7. augescens. From the Mexican and Central American 7. puberula, T. augescens differs in its more coriaceous texture, the longer and narrower lateral pinnae, and the usually distinct terminal ‘‘pinna.’”’ The costal scales of T. augescens are longer, more numerous, and of lighter color than those of 7’. puberula. The geographic ranges of T. augescens and T. puberula do not overlap. Thelypteris augescens hybridizes with T. normalis in Cuba and Dade Co., Florida. Hybrids also occur with T. ovata var. ovata in 30 AMERICAN FERN JOURNAL southern Florida and the Bahamas. In. both cases the hybrids are sterile and the spores abortive. 6. THELYPTERIS ovata R. St. John in Small, Ferns Southeastern States 239, cum tab. 1938. Dryopteris normalis var. harperi C. Chr. Dansk. Vid. Selsk. Skr. VII. 10: 182. 1913. Hotoryre: Hawkinsville, Pulaski Co., Georgia, R. M. Harper 1382 (US! photograph UC!; isotypes MO! NY! photograph UC)). Hoxorype: Lecanto, Citrus Co., Florida, R. St. John 394 (NY}). Hasitar anp Distripution. Hammocks. and limestone sinks along the southeastern coastal plain from 8. Carolina to Alabama Andros and New Providence in the Bahamas. 6a THELYPTERIS OVATA VAR. lindheimeri (C. Chr.) A. Reid Smith, comb. nov. Dryopteris normalis var. Lindheimer C..Chr. Dansk. Vid. Selsk. Skr. VII, 10: 182. 1913. Hotorype: Texas, F. Lindheimer 742 (B! photograph UC!; isotypes MO! UC! US! photograph UC)). Thelypteris X lindheimeri (C. Chr.) Wherry, Southern Fern Guide 346. 1964. Hapirar anv Disrripution: Riverbanks and moist canyons from central Texas south to northern Veracruz and Puebla; apparently most common in Texas and the states of Nuevo Leon, Tamaulipas, and San Luis Potosi in Mexico. The characters which best distinguish 7. ovata from its closest relatives, namely T. normalis and T. augescens, have already been discussed. Two forms of 7’. ovata var. ovata occur in southern Florida which are rather different from each other, and at the same time are unlike specimens from central and northern Florida. One of these is found chiefly in the Everglades and is characterized by fronds frequently greater than one meter long and pinnae 18-25 em long. The second form occurs in lime sinks and along roadsides and river banks. Its fronds are seldom greater than 0.5 m long and its THELYPTERIS NORMALIS COMPLEX ot pinnae are much shorter, often less than 10 em long. These two forms are alike in nearly all other characters. When grown under uniform greenhouse conditions, these forms show much less pro- nounced size differences and there is little doubt that they belong to the same species. The presence of linear scales (mostly 0.5-1.0 mm long) on the costae below is the best character for distinguishing var. lind- heimeri from the type variety. In addition, the blade is usually less broadly deltoid than in var. ovata. There are often minute hairs about 0.1 mm long on the lamina above in var. lindheimeri; these are absent in var. ovata. The two varieties are allopatric. Taste I. SUMMARY OF CHROMOSOME NUMBERS IN SPECIES AND HYBRIDS OF THELYPTERIS SECT. CYCLOSORUS FROM THE SOUTHEASTERN UnitTep STATES AND NEIGHBORING REGIONS.! Chromosomes populations Taxon at meiosis sampled T. augescens 72 II 2 T. dentata 72 II 3 T. normalis 72 II 28 T. ovata var. ovata 36 II 13 T. pate 72 1 ey chaidtadpilact var. versicolor 9 T. augescens X normalis ca. 36 II + 721 2 T. augescens X oyata ca. 36 II + 361 2 T. normalis X ovata ca. 36 II + 361 4 ‘For most populations, several plants were counted. LITERATURE CITED ie laa C. 1913. A monograph of the genus Dryopteris. Part I. The opical American pinnatifid-bipinnatifid species. Dansk. Vid. Selsk. er ., Ser. VII. Nat. og Math. Afd. 10: 55-282. Morton, C. V. I 967. Studies of fern types. I. Contr. U. 8. Nat. Herb. 38: Sma, J. K. 1938. Ferns of the Southeastern States. Science Press, Lancaster, F 32 AMERICAN FERN JOURNAL Srroruer, J. L., and A. R. Smirx. 1970. Chorology, collection dates, and taxonomic responsibility. Taxon 19: 871-874. Wuerry, E. T. 1943. Observations on Florida ferns. Amer. Fern J. 32: 139-145. —__—_——. 1964. The Southern Fern Guide. Southeastern and South-Midland United States. Doubleday, Garden City, New York. HERBARIUM, DEPARTMENT OF Borany, UNIVERSITY OF CALI- FORNIA, BERKELEY, CALIFORNIA 94720. Asplenium pinnatifidum x trichomanes—A New Record for Indiana GERALD J. GASTONY One of the most interesting aspects of the 1970 foray of the American Fern Society was the discovery of a new record for the State of Indiana, the rare hybrid Asplenium pinnatifidum X trichomanes. The population dynamics demonstrated by Wagner (1954) in the Appalachian Spleenworts are again illustrated by this natural hybrid, which was previously known only as a single plant from southern Illinois (Wagner & Wagner, 1969). The specimen here reported was discovered by the author—and nearly simultaneously by Rolla M. Tryon—in a pocket on a south-facing sandstone cliff at McBride’s Bluff along the east fork of White River in Martin county, Indiana, about six miles north of Shoals on August 22, 1970. Rolla Tryon and the author had been attracted to this particular portion of the area’s extensive sandstone cliffs by Amy and James Mason and Lynda Cole, other foray members, who were investigating a peculiar variant of Woodsia obtusa on an adjacent east-facing spur of the outcrop. Like the specimen reported by Wagner and Wagner, the present hybrid is morphologically intermediate between the parental species. The pinnae are quite like those of A. trichomanes (Plate 6), but express the influence of the A. pinnatifidum: genome in being more broadly attached to the rachis. The influence of A. pinnatifidum is also evidenced by the long-tapering lobed apex of several of the hybrid’s leaves. The petiole and proximal part of AMERICAN FERN JOURNAL VoLuME 71, PLATE 6 ASPLENIUM SPECIMENS FROM SANDSTONE CLIFFS aT McBRripe’s BLuFF NEAR SHOALS, INDIANA. Fig. 1. REPRESENTATIVE LEAF OF A. PINNATIFIDUM Nutt. Fig. 2. REPRESENTATIVE LEAF oF A. TRICHOMANES L. Fig. 3. A. PIN- NATIFIDUM X TRICHOMANES, 34 AMERICAN FERN JOURNAL the rachis is brown (with brown streaks often extending to the second to fourth pair of pinnae) and half-terete to adaxially chan- neled as in A. trichomanes, whereas distal to that the rachis is green and adaxially more flattened as in A. pinnatifidum. The minute, clathrate, trichomoid scales on the petiole and rachis of the hybrid are similar to those in A. pinnatifidum, whereas these are absent in A. trichomanes. With few exceptions, the sporangia of the hybrid are clearly abortive. Occasional sporangia in several sori are rather normally developed, but even these evidently bear aborted spores whose number is uncertain but appears to be con- siderably fewer than the 64 of the parental species. This may indicate the incipient development of a sporogenetic system co- ordinate with apogamous reproduction. Because fixatives were unavailable in the field and the specimen did not appear fresh enough for cultivation upon arrival at Indiana University, the hybrid was not investigated cytologically. Nor have the populations of A. trichomanes at McBride’s Bluff been examined to determine which of the ploidal levels known for this species may be represented. The leaves of the present specimen are morpho- logically quite similar to those figured by Wagner and Wagner in which the parental A. trichomanes population was diploid. Thus it is not morphologically apparent that A. trichomanes of higher ploidy than diploid has been involved in the parentage of this hybrid. Voucher material of this specimen, Gastony 949, has been deposited in the Indiana University Herbarium at Bloomington (IND) and at the Gray Herbarium (GH). LiTeratureE Cirep Waener, W. H. Jr. 1954. Reticulat wae Evolution 8: 103-118. Waaner, W. H. Jr. and F. S. Wacner. 1969. A new natural hybrid in the Appalachian Asplenium complex and its taxonomic significance. Brittonia 21: 178-186. in the Appalachian Aspleniums. DEPARTMENT oF Botan Y, INDIANA University, BLoomineron, _ Inprana 47401. Dwarp OstTRICH FERN 35 A Dwarf Ostrich Fern H. Lou Gipson In mid-summer 1956, I found a small, isolated, patch of ferns of a species that I did not recognize. The ferns were growing in a strip of moist, alluvial woods along the bank of the Moose River in the Adirondacks, near McKeever, Herkimer County, New York. They appeared similar to typical Matteuccia struthiopteris var. pensylvanica (Fig. 1), but had slightly more rounded an much shorter fronds (Fig. 2). They did not appear to be repro- ducing by offsets from their rhizomes, nor do I recall seeing any fertile fronds. The patch was separated by about 50 feet from Ostrich Ferns growing along the river bank. The land there was about one foot higher and somewhat drier than in the rather mucky flat where the small Ostrich Ferns grew. I brought a few plants back to my wildflower garden. They persisted successfully and did not change in character with re- spect to size, offsetting, or fertility. In 1967 one plant did produce a stunted and twisted “fertile” frond about one inch long, tight to the crown. But no prothallia could be grown from it. In 1964 I studied stem sections in an attempt to identify the species. The vascular strands of the dwarf Ostrich Ferns seemed identical to those of the regular ones. In order to exclude the possibility that soil conditions in my garden maintained an environmental dwarfing, I did a few ex- periments. Small plants were paired as to size with young offsets from large Ostrich Ferns. Three pairs were planted in various parts of the garden. In each case the regular ferns grew to a large size, while the small ones remained unchanged now for several years. Professor Wagner, who has examined the specimens, is of the opinion that the dwarf condition is genetically fixed. The dwarf plants grew equally well in wet and drier areas of the garden, whereas the normal plants attained greater heights in a wet section than in a dry one. In 1965 I returned to the natural site to study the original AMERICAN FERN JOURNAL VoLUME 71, PLATE 7 Weeatcaaa. 1 @ @ MATTEUCCIA STRUTHIOPTERIS FRONDS. Fia. 1. NORMAL FROND 60 INCHES LONG. Fig. 2. DwaRF FROND 15 INCHES LONG. SHORTER NOTES 37 patch. Trees had been cleared along the strip for road building, causing a denser undergrowth. A washed-out dam had lowered the river by about two feet. Ostrich and Cinnamon Ferns were more profuse than formerly. The area was searched, but every likely Matteuccia of small stature could be traced by the long- creeping rhizomes back to alarge, full-grown Ostrich Fern. A more detailed and lengthy search, perhaps with a ready means for identification, might reliseoeee the original dwarf population. The small ferns are probably overgrown by the normal species Some perusal of the literature and consultations with fern authorities have not uncovered any mention of a dwarf variety of the Ostrich Fern. Perhaps this report will prompt further discussion. ROcHESTER ACADEMY OF SCIENCE, RocueEsTeEr, N. Y. 14617. Shorter Notes NIPHIDIUM LONGIFOLIUM, A NecEsSARY NEw CoMBINATION.— In Copeland’s “Genera Filicum,”’ Niphidium, based on N. ameri- canum (Hook.) J. Smith (Hist. Fil. 99. 1875), is recognized as a distinct, monotypic genus, a native of the Andes of Ecuador. This species has been generally placed in Cyclophorus (now correctly Pyrrosia), which is a genus strictly confined to the Old World. Although N. americanum is somewhat like Pyrrosia because of its densely tomentose appearing undersurface, it is probably not closely allied. As Copeland pointed out, Pyrrosia has stellately branched. peltate scales (Fig. 3) on the undersurface, whereas Niphidium has (under the compound microscope) beautifully clathrate scales bordered with elongate, hyaline hairs (Fig. 1). The hairs are usually forked at the apex and two cells wide at the base (Fig. 2); each cell forms one of the forks. These hairs are so numerous and dense as to make the lower surface appear densely tomentose. But the frond scales of Niphidium are not truly peltate and stellate, and so a resemblance to Pyrrosia is strictly super- ficial. In its Anazetum-like venation and its large, round sori VouLuME 71, PLATE 8 AMERICAN FERN JOURNAL SHORTER NOTES 39 uniseriate between the prominent lateral veins Niphidium is like Pessopteris, and may be related to that, which is however essentially scaleless on the lower surface. In calling this plant Niphidium americanum (Hook.) J. Smith, Copeland overlooked an earlier specific name that is available. In his paper on Cavanilles’ fern types, Christensen indicated that Polypodium longifolium Cav., from the type specimen, is identical with N. americanum. Therefore, the following new combination is necessary: NipHipium longifolium (Cav.) Morton & Lellinger, comb. nov Polypodium longifolium Cav. Deser. 245. 1802. Type: et near Monat ea aha ee Née (MA, seen by Christens Beceggebi americanum Hook. Sp. Fil. 5: 54. 1864. Synrypes: Cuenca, Ecuador, Jesh. and Bafios, Ecuador, Spruce 5248 (both pre- sumably K). Niphodolus americanus (Hook.) Diels in Engl. & Prantl, Nat. Pflanzenfam. 25. 1899. - Cyclophors americanus (Hook.) C. Chr. Ind Fil. 198. 1905. Cyclophorus longifolius (Cav.) C. Chr. Dansk Bot. Ark. 9(3): 11. 1937. It should be pointed out that under Pyrrosia the epithet longi- folia is not available because of the quite different Old World species P, yeh ae (Burm. f.) Morton (J. Washington Acad. Sci. 36: 168. 1946)—C. V. Morton anp Davin B. LELLINGER, National Ae useum of Natural History, Washington, D. C. 20560. Bu.sous ApDER’s-TONGUE Common in Lovtstana.—Bulbous Adder’s-tongue is a very small, easily overlooked fern. I first saw it in March, 1970, accidently while looking for Oenothera spachiana seedlings for my wildflower garden. Most manuals consider this fern to be rare or at least uncommon. Brown and Correll (Fern and Fern Allies of Louisiana, 1942) listed it from four southeast parishes (East Feliciana, East Baton Rouge, Livingston, and Orleans), one southwest parish (Jefferson Davis) and two northeast parishes (Richland and Caldwell). They noted that it was common Fig. 1. SCALE FROM THE ABAXIAL SURFACE OF NIPHIDIUM LONGIFOLIUM, X 24 (Ginter 59). ele 2. SAME, DETAIL OF BASE OF DOUBLE-CELLED HAIR, X 576. Fig. 3. ScaLES FROM THE ABAXIAL SURFACE OF -PyRROSIA ACRO- STICHOIDES, X 50 nse ten 18054) 40 AMERICAN FERN JOURNAL only on the Copenhagen Prairie of Caldwell Parish. The University of Southwestern Louisiana has specimens from Bienville, Winn, Caldwell, and Rapides Parishes. Louisiana Technological Univer- sity has a specimen from Union Parish. The small number of specimens and the information on the labels indicate that most’ collectors consider this fern to be rare. From mid-March to mid-May, 1970, I made field trips looking specifically for Ophioglossum crotalophoroides. I have collected the fern in the following twenty-six parishes: Allen, Beauregard, Bienville, Bossier, Caddo, Caldwell, Catahoula, Claiborne, DeSoto, Evangeline, Franklin, Grant, Jackson, LaSalle, Lincoln, More- house, Natchitoches, Ouachita, Red River, Richland, Sabine, Union, Vernon, Webster, West Carroll, and Winn. Other parishes could have been added had I not been restricted in my collecting by a full teaching load. The specimens referred to above raise the number of parishes where this fern is known to grow to thirty-two. Ophioglossum crotalophoroides is apparently absent from four parishes I checked: East Carroll, Madison, Tensas, and Concordia. They lie along the Mississippi River, and I could not find any sandy soil in them. No plants were found on the clay soils character- istic of these parishes. All collections were from sandy soils of cemetaries, pastures, road banks, power lines, pipe lines, etc. Typically these sites are mowed during the summer. In such habitats the plants of Bulbous Adder’s-tongue are extremely nu- merous. Some cemetaries I visited contained thousands of these plants. I think there are almost as many (or as many) Ophioglossum crotalophoroides. plants in Louisiana as there are of such common ferns as Christmas Fern and Bracken; it simply has been over- looked by collectors. One possible explanation of this is that during late March, when O. crotalophoroides is at its peak, there are no common Showy associates—except possibly the tiny bluets, Hous- fonia pusilla, H. minima, and H. pygmaea). At this time few botanists bother to éollect in such “unfruitful” territory. Also, one can walk over those ferns ten times and fail to see them. But this small fern can be seen with any degree of regularity only from the prone position. I suggest that collectors look specifically in the RECENT FERN LITERATURE 41 proper habitats for this fern and that they get down to ground level where the plants are. Ophioglossum crotalophoroides is much more common than previously supposed.— Dr. R. DALE THoMAs, Department of Biology, Northeast Louisiana University, Monroe, La. 1201 Recent Fern Literature FLORE DE LA NovuvELLE-CALEDONIE ET DEPENDANCES, NO. 3, Pr¥ripopnytes, by G. Brownlie. Published by Muséum N atidina d’ Histoire oe 16 rue Buffon, Paris V, France, pp. 1-307. t. I-XXXIX. 9:—This beautifully prepared book is the first fern flora of any — the South Sea Islands for many years. Many of the ferns of New Caledonia are endemic, and extremely inter- esting because of their variability, in which they are comparable to the ferns of Hawaii. . Brownlie’s treatment has full keys, descriptions, and citation of synonyms, misapplied names, and specimens. The work owes much to Madame Tardieu-Blot, who translated the descriptions into French and who had the excellent drawings prepared in Paris. In fact, the whole style and format is exactly like one of Madame Tardieu’s own books on the fern floras of Gabon and. Cameroun; however, Madame Tardieu disclaims any responsi- bility for the taxonomic treatment or other details. The comments that I give below are on minor matters. They do not detract from the value of this excellent book, which should be in the hands of -all fern students. Brownlie splits up the traditional family Polypodiaceae into 13 families, following Alston (cf. Taxon 5: 23-25. 1956), who had a much better treatment in my opinion than that of Ching, Cope- land, or other more recent authors. However, one runs into difficulties in trying to describe and key out these families, and Brownlie’s key will not work well for some genera. Grammitidaceae has as one of the key characters that the petiole is generally articulate, but this is never true in this family. Vittariaceae is keyed as having the sori marginal or submarginal, but they are strictly dorsal in Antrophyum. Acrostichum, Syngramma, and 42 AMERICAN FERN JOURNAL A spleniopsis, which have the sori strictly dorsal, are keyed solely under the heading of sori marginal or submarginal. This merely points out that it is impossible to key out the families as recognized by single key characters. Probably the only way is to do as Cope- land did in his “Genera Filicum,” and have Aspidiaceae appear in ten different places in the key to the families, Pteridaceae in nine places, and so forth. Such keys are essentially keys to genera or groups of genera, rather than to families. Many of the family names used are suspect or definitely in- correct. Azollaceae is, for instance, attributed to Christensen in Verdoorn (1938), but at the place cited there is no Latin diagnosis. Presl called his “Lindsaeaceae” a “Sectio,” and since a section is a group below the rank of a genus, it can not be a proper category for a family name. The Code indicates that a family name must be called either a family or an “Ordo” (Art. 18, paragraph 3). Vittari- aceae was called by Presl a “Tribus,” and a tribe is also an in- admissible category for a family name. Psilotaceae is attributed to Payer (1850), but Payer had it in French as “Psilotées” ; since botanical nomenclature must be in Latin, names published origi- nally in French are not valid; ef. Code, 1966 edition, p. 213, where “Casuarinées” Mirbel, 1810, is rejected in favor of Casuarinaceae R. Brown, 181421 Many stated types are in reality lectotypes only (e.g. Diplazium echinatum C. Chr.). Vieillard 1520 from Canala can not be the 1s Impossible, since this was not one of the species originally included in the genus. The type of Lastreopsis is Polypodium recedens J. Smith, not L. tenera (R. Brown) Tindale, which may be 1 This book is remarkably free of t hi i i. ypographical errors; one noted is the citation of Athyriaceae Alston as “Taxon 3:36. 1936’’; it should be 5: 25. 1956. RECENT FERN LITERATURE 43 a taxonomic synonym but which is based on an entirely different type specimen. The only genus in which the treatment is unsatisfying is that of A splenium, in which a number of species and varieties described from New Caledonia are placed in a table showing their possible origin as hybrids, but with absolutely no diseussion of these, or any citations, descriptions, or typifications. Some of the compound epithets are spelled without a hyphen, e.g. Hymenophyllum le ratii and Adiantum capillus veneris, but the Code requires such specific names to be hyphenated. The specimens that are referred to Lycopodium nutans Brack. (1854) are probably L. proliferum Blume (from unpublished annotations by Adelbert), and if this is correct the Blume name has priority. Cyathea propinqua Mett., a Fijian species of the subg. Sphaero- pleris, is cited as a synonym of C. vieillardii Mett., which belongs in subg. Cyathea, according to Holttum’s classification (Blumea 12: 272. 1964). Asplenium adiantoides (L.) C. Chr. (1905) is accepted as a correct name, but it is an illegitimate later homonym of A. adiantoides Lam. (1786); I believe that the correct name is A. polyodon Forst. Thelypteris uliginosa (Kunze) Ching has Cheil- anthes setigera Blume (1828) cited as a synonym; T’. uliginosa is a different species fron the Javan T. setigera; the correct name under - ‘Thelypteris is T. torresiana (Gaud.) Alston, which has recently been ge Macrothelypteris torresiana (Gaud.) Ching (cf. Holttum, Blumea 17: 27. 1969). Marattia rolandi-principis is described as being cme pinnate, but it is sometimes bipinnate; Rosenstock distributed material as M. rolandi-principis var. subbipinnata Rosenst., an unpublished name, and the bipinnate plants were collected by Guillaumin and Baumann (no. 8663). Nephrolepis floccigera (Blume) Moore is cited as a synonym of N. cordifolia, but from the holotype (from Celebes) it is by no means that, for it has the pinnae about 12 cm long or more, not about 2 cm long as in N. cordifolia. Blechnum vieillardii var. simplex Fourn. is a nomen nudum at the place cited. There are few omissions. The genus Belvisia [Hymenolepis] ‘perhaps needs further study; only B. mucronata is reported. 44 AMERICAN FERN JOURNAL Several collections of Hymenolepis revoluta var. planiuscula (Mett.) Hieron were cited from New Caledonia by Christensen (Dansk Bot. Ark. 6(3): 59. 1929). Cephalomanes australicum v. d. Bosch was described from the Isle of Pines, which is included within this New Caledonian flora. Pyrrosia varia (KKaulf.) Farwell has been reported from New Caledonia but is not accounted for here. Otherwise, the only omissions noted are: Blechnum trregulare Carr. (New Caledonia, MacGillivray F25, holotype BM, Morton photograph 6685), B. diversifolium var. paleaceo-selosum Rosenst. (Fedde Repert. 8: 75. 1910), Lomaria attenuata f. monstrosa Compton, L. lenormandii f. aprica Compton, Asplenium adian- toides var. tripinnatum Compton (= A. robustum Blume?), Cyathea albifrons var. lata Compton, Hymenophyllum mnioides f. amplior Compton, and Lycopodium squarrosum var. pacificum Compton.—C.V.M. Tue Ferns AND FERN ALLIES OF THE SIERRA NEVADA IN CaLiFoRNIA AND Nevapa, by John Thomas Howell. The Four Seasons 3(3): 2-18. 1970.—In anticipation of his forthcoming “Flora of the Sierra Nevada,” Howell has presented here a list of the ferns and fern allies with a key and an indication of the north- ernmost and southernmost known stations. There are altogether 62 species treated, one of which is introduced (Cyrtomium falca- tum) and several of which are probable hybrids. ‘“The Four Seasons” is a relatively unknown periodical issued quarterly and consists chiefly of taxonomic notes on Californian plants. It is published by the Regional Parks Botanical Garden, Berkeley, Calif. 94708; the subscription price is $6.00 per volume.—C.V.M. American Fern Society Report of the 1970 Fern Foray Southern Indiana sandstone cliffs and wooded ravines, the richest sites for ferns in the state, were the focus of the Fern Foray, August 20-22, 1970. The first stop was near Taswell, in the Hoosier National Forest, an unusual deciduous forest having elements characteristic of coniferous woods, such as Hemlock, Yew, Moun- AMERICAN FERN Society 45 tain Laurel, and Partridge Berry. There were abundant plants of Asplenium platyneuron growing in open, grassy sites leading up to the woods. Pteridium aquilinum var. latiusculum, and Equisetum hyemale also occurred in these open areas. One of the Gentians (Sabatia angularis), the False Foxglove (Aureolaria flava), and many composites were in good bloom. In the cool ravines were many plants of Botrychium virginianum and B. dissectum f. obliquum, Adiantum pedatum, Diplazium pycnocarpon, D. acrosti- choides, Athyrium Filizx-femina var. asplenioides, and Thelypteris hexagonoptera. Several orchids, the showy Orchis spectabilis, and Rattlesnake Plantain (Goodyera repens), as well as Bloodroot, Hepatica, Baneberry, Anemone, and Trilliums were evidence of the rich Spring flora in these woods. On the higher slopes of the ravines, among rocks fragmented from the cliffs, there were numerous plants of Polystichum acrostichoides, Dryopteris margi- nalis, and fewer of D. spinulosa var. intermedia and Cystopteris fragilis. On the spectacular, sheer cliffs, ledges were covered with Polypodium virginianum, and a few leaves of P. polypodioides were found. The pocked faces of the sandstone, especially where mosses were established, provided pockets for Asplenium pinnatifidum and A. trichomanes. One of the rare Saxifrages, Sullivantia Sulli- vantii, kept company with these Aspleniums and Camptosorus rhizophyllus. Some of the more energetic climbers who scaled the cliffs found, on the upper slopes, one of the uncommon ferns of Indiana, Dennstaedtia punctilobula. On the afternoon trip to Pioneer Mother’s Memorial Forest, near Paoli, many ferns of the ravine were seen again. This woods is one of the few remaining stands of primitive hardwoods in the state, with fine specimen trees of White Oak, Walnut, and Tulip Popular. There were several unusual orchids here—Lily Tway- blade (Liparis liliifolia), Puttyroot (Aplectrum hyemale), and flowering specimens of the Nodding Pogonia (Triphora triantho- phora). During the evening program, at our motel, in Jasper, Rolla Tryon reviewed the recent glacial history and vegetation of the region, and discussed the species that were collected during the VoutuME 71, PLATE 9 AMERICAN FERN JOURNAL STANDING LEFT TO RIGHT: NEILL Haun, James MonTGONERY, JEANETTE Oxtver, Wituiam anp GARNET Meinpers, BENTON Strpp, Guedory ANDER- son, Donatp Hu TTLESTON, JOHN CLEMENTS, SUSAN AND G ERALD GASTONY, OBERT LOMMASSON, Meise Faust, Roa TRYON, James. MAYSILLEs, Mrs. R. BENEDICT, VicTOR Rigs, Sow maeta Drow, W. AND Mrs. Rovas Eva Sopot; Seatep: Hixases Joe Hosuizakt, JEANETTE Gross, Exivor Henry, Lynpa Coie, LeRoy Henry, Auice TRYON, Amy Mason, Berry Ditton, Rateu Besmnicr. PHOTO TAKEN BY STEVE Wikinane: AMERICAN FERN SocreTy 47 day. Neill Hall reported on the activities of the spore exchange and requested spores of any species, to replenish older supplies. Donald Huttleston spoke of the living fern collections at Long- wood Gardens, which is actively acquiring living collections from all parts of the world. Mrs. LeRoy Henry showed her colored movies and commented on the 1967 Fern Foray, in Texas, and the 1969 International Botanical Congress, at Seattle, including some of the attractive gardens of the area. The film had many good views of Fern Society members as well as habitats in Texas ana Washington. The second day’s trip was to MacBride’s Bluff, another sand- stone cliff area, near Shoals. Woodsia obtusa was abundant on the open road banks growing with the blue-flowered Lobelia siphilitica. Great colonies of Polypodium polypodioides and Camptosorus rhizophyllus covered boulders below the cliffs. A few plants of Cystopteris bulbifera were found on the cliffs and one of the un- common ferns of Indiana, Chezlanthes lanosa, occurred on pre- carious ledges. Plants of Asplenium pinnatifidum and A. tricho- manes grew in association in pockets on the cliff faces. The rare hybrid between them, a fern new to the state, was a most appro- priate find by Gerald Gastony, a new oe on the staff at Indiana University. An extra stop was made at the Martin State Forest, near Shoals, for several of the group who wanted to see Dryopteris goldiana. We were especially pleased to have among the thirty-three members and guests attending the Foray, several Hoosiers, enthusiastic and knowledgeable about their flora including, Gregory Anderson, John Clements, Lynda Cole, Amy and James Mason, Jeanette Oliver, and especially James Maysilles, of Han- over College, who arranged for our comfortable lodging and a fine picnic lunch. We are much appre¢iative of his help and that of Rolla Tryon in guiding us to some of their favorite haunts in southern Indiana.—A.icE F. Tryon, Gray Herbarium, Harvard University, Cambridge, Mass. 02138. Statement of Ownership, Management and Circulation In accordance with the rules and regulations of the United States Post Office, as established under the act of October 23, 1962, Section 4369, Title 39, United States Code, the following statements are published Title: AMERICAN FERN JOURNAL fis sac ee Issue: Quarterly ea Neamt March 31, June 30, mber 30, and Decem 1) Location oe ‘Office of Publication “Printers: 3110 Elm Avenue, altimore, Maryland 21211 Location of Bosiness Office of Publishers (Not Printers): Dr. LeRoy s crete Section of Plants, Carnegie Museum, Pittsburg, 1521 Publisher: fame Fern Society, Inc., Section of Plants, Carnegie um, Pittsburg, Pa. 15213 Editor: De | David B. Lellinger, ee . agg Smith- sonian a Washington, D. C. 2 Monae Editor: Non Owner: — ha Socrery, Inc., Section of Plants, Carnegie Museum, Pittsburgh, Pa. 1521 3 Bondholders, Moraceae and other Security Holders: None Average No. each Single issue issue during pre- nearest to ceding 12 months filing date Total number of copies printed: 1400 1400 Paid circulation : Sales through dealers, etc. 0 0 2. Mail subscriptions 951 945 Total paid circulation 951 945 Free recite imi samples 0 0 Total distribut 951 945 Office use, betover: ete. 449 455 Total 1400 1400 The statements made above are certified to be correct, and are signed by: aie B. es editor-in-chief of the American Fern Journal on Septem- ber 28, 1970. Exotic and Hardy Ferns 3 Begonias BOLDUC'S GREENHILL NURSERY Lo 2131 Vallejo Street St. Helena, California 94574 Open Saturdays and Sundays from 10 A.M..to 4 P.M. and by appointment Phone $63-2998—Area Code 707 | _-... Mall orders accepted TRIARCH INCORPORATED -~A Service to Biologists = To better serve the biological community: pathology, bacteriology, general zoology, vertebrate histology, vertebrate embryology, and parasitology. _2. TRIARCH offers to prepare slides of new or spe- cial items for a nominal fee or in exchange for pre- served materials, 3. TRIARCH advertises in biological journals to help support the financial needs of the associated So- ciety. For our current catalog No. 15, or to request special service, write to: Paul L. Conant, President Ripen, Wiseensin 54971 Vou. 61 Aprit-JUNE, 1971 No. 2 American Fern Journal A QUARTERLY DEVOTED TO FERNS Published by the AMERICAN FERN SOCIETY ts DAVID B, LELLINGER C. V. MORTON ROLLA M, TRYON IRA L, WIGGINS CONTENTS Lindsaea (Schizoloma) ensifolia Swartz in Hawaii.W. H. Wacnsr, Jr. The Fern Collections in Some European Herbaria, VII C. V. Morton 59 An Appendageless Psilotum. Introduction to Aerial Shoot Morphology Ausert S. Rovrra 75 Vascularization of Fern Leaves Rozert C. Lowmasson anv C. H. Youne, Jr. 87 Shorter Note: A Wrongly Localized Species of Pyrrosia........-.---- 94 95 Recent Fern Literature..........-. ee ee ee ee Missour! BoTANIcAL Jup 16 1971 GARDEN LIBRARY Che American Fern Society €ouncil for 1971 Warren II. Wacner, Jz., Department of Botany, University of — Ann Arbor, Michigan 48104. Presiden Joun T. Micxer, New York Botanical Garden, Bronx Park, rats New York. 1 Vice-President Ricuarp L. "ae. , Department of Botany, University of Rode Tieng, ——— Rhode Island 02 tary LeRoy K. Bgteg Division of Plants, Carnegie Museum, Plier eae poleania 213. Gower - Davin B. Taccecce. Smithsonian Institution, Washington, D. C. Editor- in Cntel —— Society Representatives Warren H. Waaner, Jr., University ea ae = = £ S. Representative Rona M. Teor, rx Harvard Unive B.S. Representative American ae Journal EDITORS Davip B. aoa eed Smithsonian Institution, Washington, D. C. 20560. C. V1 MIGSTON 5... : Smithsonian Institution, Washington, D. C. 20560. Roa M. Taro, = Gray Herbarium, Harvard University, Cambridge, Mass Tra 1, Wioarns... ae Herbarium, Stanford Gsivernity 3 Stanford, “Calif An illustrated A say Pag oted to the general 10) Him Ave — owned b gy at _Aghd and pubieget at 3110 * Baltimore, Md. 21211. Second cam pee age paid at Baltimore. aioe ‘of the Jour- nal are open db enon a who wish to arrange exc poi Sa ion membership lists assist specimens from different localities Matter for a n m shoul be addressed to to the Edito oe Changes , applications for membership, su iptions, orders for back pcan igre okt be 0 rp ena to the ateaeeei. 2 ns why mitts rod fee $0. sail sent free to mem- the American Fern Society (annual dues, $5.00; sustaining membership, ae: life membership, $100.00 A txcencted ta reprints, if ordered in advance, il be furnished at cost, to Postage. Pigs $5.00 to —. each; single back ae: is onive Cumulative eg pons each; over 80 pages, $2.50 mi to ne! Soni -25, 50 cents. Ten percent discount on sare of six volumes Library and Berbarium Dr. W. H. Wagner, Jr., Department of Botany, University of Michigan, Ann Arbor, Mic 64, is Librarian and Curator. oh aa Sg y borro books and specimens at any time, the borrower paying all shipping seates Spore Exchange Mr, Neill D. Hall, 1225 Northeast 95th Street, oeeiged —— 98115, is Director. Spores PPR Se eh eee a a to th oat eRe reine sac ob Bp IMPORTANT ERRATUM Due to an error on the part of the printer, the January-March, 1971, issue of the American Fern Journal was indicated at the top of the front cover, at the top of page 1, and at the right hand top corner of the plates, as being “Vol. 71.” The correct volume num- ber is 61. Please correct your copy in order to avoid problems of bibliographic citation in the future. American Fern Journal Vou. 61 APRIL-JUNE, 1971 No. 2 Lindsaea (Schizoloma) ensifolia Swartz in Hawaii W.H. Waaner, Jr.! The ferns and other pteridophytes of the Hawaiian archipelago include approximately 180 species, of which 20 were naturalized since 1850, having been introduced by man (Wagner, 1948, 1 The islands have received an enormous amount of exploration by botanists over the years, but discoveries are still being made. Today we tend to assume that all of the major indigenous genera and species groups have already been found. Nevertheless, I wish to report a wholly distinctive species of fern not heretofore col- lected in the islands. As will be described below, the plant in question grows in a rather unlikely habitat in association with other species which more or less resemble it.? The fern could be readily overlooked; in fact, it was almost an accident that we noticed it at all. On December 16, 1969, a group of botanists and ecologists visited the Puhimau “Hot Spot,” an area along the Chain-of- Craters Road in Hawaii Volcanoes National Park, just south of ' Research supported by NSF Project GB-8113, The Evolutionary Charac- ters of Ferns. ia ab to acknowledge the aid of Charles Cooper, J. Linsley Gressitt Ne U. Kramer, D. B. Lellinger, D. Mueller- Dasnboks, and Garrett mat. * One es is Pteris vittata L. Although adult plants of a species are obviously ferent from Lindsaea ensifolia, the juveniles are m re difficult to distinguis hey differ in having ir pinnae short to fe eee are ternate and perhaps even simple at early stages, and become pinnate only at later stages—D.B.L. Volume 61, No. 1, of the JourNaL, pp. 1-48, was issued March 23, 1971. ~ AMERICAN FERN JOURNAL Vo.tumE 61, PLate It 2 FuMEROLE AREA IN Hawan VoLcaNogs N \PPE 2INCE OF VEGF’ aces : APPE \RANCE OF VEGETATION AND PLUMES OF HOT STEAM. Fic. 2. OBSERVER STANDING NEAR THE EDGE OF A LARGE STEAM VENT ATIONAL ParRK. Fic. 1. BLEAK LINDSAEA ENSIFOLIA IN Hawall 51 Puhimau Crater (briefly described by Doty and Mueller-Dombois, 1966, pp. 322-323). Surface and steam vent temperatures here run from 48°-89° C, and the ground is warm to the touch. The roots of many of the plants, including the conspicuous grass Andropogon virginicus and scattered small trees and shrubs o Metrosideros are only at the surface and do not penetrate into the t earth. For this reason the plants are not anchored and can be literally moved around and re-arranged like pieces of furniture on the white, ash-like soil. Plumes of steam arise in an eerie fashion in this other-worldly habitat (Fig. 1, 2). Around the steam vents are dense growths of mosses and ferns (Plate 11). The ferns are mainly Swordferns, Nephrolepis evaltata, but occasional patches of Ophioglossum nudicaule occur among the mosses. The Swordferns tend to become progressively smaller the closer they grow to the steam, as shown in the lower half of Plate 11. While we were examining the Swordferns around the steam vents we first noticed small plants that seemed especially in texture to be different from the juvenile plants of other fern species known to occur in the Kilauea area. Careful examination of these plants and their associates finally revealed older stages of this fern, which we realized was a species unreported in the islands. Colonies of it, usually vastly outnumbered by the associated and grossly similar-appearing Nephrolepis, were later located around several of the steam vents. Subsequently the plant was identified as Lindsaea (Schizoloma) ensifilia, about 2600 miles from its nearest other localities in Samoa. In the following paragraphs comments will be made upon the leaf structure, juvenile and mature, and the significance of the occurrence of this fern in Hawaii. The early heteroblastic stages (Plate 12) are different in appear- ance and structure from the mature stages in Plate 13. The earlier stages look similar to the corresponding juvenile stages of the Tree- fern Cibotium. But the little plants of Lindsaea are glabrous and much thicker-textured than those of Cibotium. The young stages have jagged, more or less lobed pinna margins with up to two pairs of pinnae. The veins are free, except the larger specimens VoLumE 61, PLATE 11 AMERICAN FERN JOURNAL ATE FORMS OF > v DEPAUPEI RESSIVELY MORE ROG UMEROLE WITH P DGE OF F THE DomiNANT NEPHROLEPIS EXALTATA (Photograph by Mueller-Dombois). E ~ “4 LINDSAEA ENSIFOLIA IN Hawall 53 (on the right side on Plate 12) which have more or less casual anasto- moses. These fronds are transitional to the mature forms shown in Plate 13. Holttum (1954, p. 342) in his discussion of this species and its relatives comments that: The main evolutionary trend in Schizoloma [a taxon that he upholds as a genus distinct from Lindsaea] is towards a simply pinnate (or even simple) frond with reticulate veins and a long fusion sorus. On this view, the common Malayan S. ensifolia represents an ad- vanced type... It is interesting to note, therefore, that the heteroblastic series of Lindsaea ensifolia leaves from Hawaii seem to follow approximately olttum’s concept of the evolutionary trend in frond evolution, a possible case of recapitulation. The largest fronds of L. ensifolia that we were able to find are consistently trifoliolate (Plate 13). Their veins are copiously netted (for details of pattern see Wagner, 1952, fig. 24,c). The leaf margins of the mature fronds are nearly entire. We were able to find only a few fertile fronds, and these were only incompletely fertile, with the sori interrupted and widely separated. Efforts should be made to obtain more fertile fronds in order to determine whether the Hawaiian form has consistently discrete sori. Mature fronds were sent to Dr. K. U. Kramer of the State University of Utrecht, Netherlands, who has made extensive studies of lindsaeoid ferns in various parts of the world. In com- menting on these specimens (letter, March 4, 1970), he wrote: It is indeed strange that only trifoliolate leaves of L. ensifolia have been found at the Hawaiian fumeroles. Offhand, without knowing the exact local situation, I should say that apparently the plants fail to reach a more mature stage with multijugate lamina. In what I take to have been poor habitats in the mountains of e.g. New Guinea even simple forms have been col- lected. In the genus Lindsaea it is not uncommon to find fertile leaves even in small, evidently far from full-grown leaves. /OLUME 6 ATE 12 AMERICAN FERN JOURNAL VouuME 61, PuaTE 1 suqsegerniay sigan HETEROBLASTIC LEAF STaGes or LinpS\EA ENSIFOLIA FROM STEAM VENT EpGes with Earner Sraces at Lert. vr Gr LINDSAEA ENSIFOLIA IN Hawall In another communication (January 7, 1970), Dr. Kramer stated that: The most intriguing thing is perhaps that this is clearly ssp. ensifolia which has a western distribution (East to Micronesia and the Solomon Islands), not ssp. agatii which I have seen from Micronesia, Mela- nesia, and Polynesia (Fiji, Samoa, Tonga), which is the eastern subspecies. [For details of the contrasts, see Kramer, 1967a, p. 579]. As Lindsaea ensifolia was not encountered in Hawaii until December, 1969, one might argue that it had been introduced by man, either intentionally or unintentionally. I am, however, hesitant to accept this argument. This is not a plant likely to be carried by man. Unlike various naturalized ferns of Hawaii (cf. Wagner, 1948), L. ensifclia is a relatively unattractive fern and is not grown at all in horticulture, to my knowledge. The area where it occurs in the fumeroles has been studied in some detail by botanists, it is true, but the appearance of the plant is such that it can be casually confused with imperfectly developed stages of other ferns, e.g., Cibotium or Nephrolepis. It can be overlooked readily in a tangle of dwarfed Nephrolepis fronds (Plate 11). The geographical separation of the Hawaiian plants from other Poly- nesian localities is very large (the nearest are Samoa, Fiji, and Rotuma, according to St. John, 1954), but such large disjunctions across the open seas are not uncommon among Pacific ferns. In fact, the geographical distribution is similar to the two other lindsaeoid ferns known in Hawaii, Lindsaea repens and Sphenomeris chinensis. Fern spores are capable of long-distance dispersal because of their extremely small size. Current research suggests that a single spore has the potentiality of initiating a new colony of ferns by gametophytic self-fertilization. Fern gametophytes tend to be self-compatible according to the recent studies of Klekowski and Baker (1966). Physically the habitat in which we found Lindsaea enstfolia in Hawaii is peculiar. However, L. ensifolia in other parts of its range is very commonly a sun fern, growing in open, lateritic savannas, AMERICAN FERN JOURNAL VouumE 61, PLATE 138 Mature Fronps or Linpsaga ENSIFOLIA FROM THE STEAM VENTS, CARE- se, ee =, Tim r VOC TNT g 'f FULLY SPREAD OUT IN PRESSING (LEFT) \ND PresseD Directity WirHoutT OPREADING (RIGHT). LINDSAEA ENSIFOLIA IN Hawall 57 and capable of withstanding the heat of direct sunlight. Holttum (1954, p. 243) states that it “has the power to live in poor soil, in exposed as well as shaded places.” In the Kilauea fumeroles it is associated with Ophioglossum nudicaule, a common associate also in the savannas of Micronesia (e.g., Guam) and Melanesia (e.g., Admiralty Islands). The very brief list of lindsaeoid ferns of Hawaii has now been expanded from two to three. These are morphologically different from one another, but they do share rather similar, broad ranges. Lindsaea repens (Bory) Thwaites belongs to subg. and sect. Odontoloma according to Kramer, and is interpreted by him (1967b, pp. 568, 569) as being ‘“‘very variable, and a number of more or less limited infraspecific categories can be distinguished.” He interprets the Hawaiian plant as var. macraeana (Hook. & Arnott) Kramer. Lindsaea ensifolia belongs to subg. Lindsaea sect. Schizoloma according to Kramer (1967a). Sphenomeris chinensis (L.) Maxon was previously designated by most recent writers as S. chusana (L.) Copel., but the latter name is evidently incorrect (Kramer, 1967a; Fosberg, 1969). All three of the Hawaiian linds- aeoid ferns range as far west as tropical Asia. Further field studies of the fumeroles of the ISilauea area are recommended to determine whether other still unreported fern species have been overlooked in these seemingly unlikely fern habitats. Outside of Hawaii, I am aware of only one previous re- port of a strongly disjunct fern population around hot effluents. T. M. GC. Taylor (1963) reported an unusual occurrence of the southern maidenhair fern, Adiantum capillus-venerts L., as follows: In British Columbia it is known from the runnels of the hot springs at Fairmont, north of Columbia Lake. Despite the fact that the nearest known colony is some hundreds of miles to the south, there is very strong i tantial evidence to support the view that it is in- digenous at Fairmont and not an escape from culti- vation. In the Hawaiian fumeroles there is no reason to believe that the disjunct ferns reported here are not also indigenous. Now that 5S AMERICAN FERN JOURNAL we know that both Ophioglossum nudicaule and Lindsaea ensifolia are found in the steam vents of Kilauea and are apparently limited to them, interest in the investigation of these unusual habitats should be stimulated. LITERATURE CITED Dory, M. 8. and D. Mue.ieR-Domsors. 1966. Atlas for Bioecology Hoes in Hawaii Volcanoes National Park. Hawaii Bot. Sci. Pa per 2. ii + 507 pp. Fiieac. a R. 1969. The nomenclature of Sphenomeris chinensis [Filic.]. xon 18: 596-598. poe er A E. 1954. Ferns of Malaya. Gov’t. Printing Office, Singapore. 643 pp. Kuaxows E. J., Jk. and H. G. Baker. 1966. Evolutionary significance of platy i in the Pteridophyta. Science 153: 305-307. KRAMER, "K. U. 1967a. The lindsaeoid ferns of the Old World. I. New Cale- donia. Acta Bot. Neerl. 15: 562-584. - 1967b. The lindsaeoid ferns of the Old World. II-III. Notes on Lindeles and Sphenomeris in the Flora Malesiana Area. Blumea 15: 545-576. Sr. eo es 1954. Ferns of potas Island, a descriptive manual. Occ. pers Bishop Mus. 21: 161-208. TayLor, ie M. C. 1963. The Ferns and Fern-allies of Firitish Columbia. okie Columbia Prov. Mus. Dept. Recreation and Conservation Handbook No. 12. 172 Waener, W. H., Jr. 1948. Ferns ontarutiead 3 in Hawaii. Occ. Papers Bishop. Mus. 20: 95-121. - 1952. The fern genus Diellia. Univ. Calif. Publ. Bot. 26: 1-212. . 1963. Pteridology in Hawaii. Hawatian Bot. Soc. Newsletter 2: 117-123. Matruaret Boranican GaRDENS, UNrversity OF MICHIGAN, ANN Arsor, Micuican 48105. FERNS IN EvuROPEAN HERBARIA 59 The Fern Collections in Some European Herbaria, VII C. V. Morton BRUSSELS There were doubtless some herb gardens in Brussels from very early times, but the first formal association of gardeners was that of the Confrérie de Sainte-Dorothée formed in 1650 and named in honor of Saint Dorothy, supposed for some reason to be the patron saint of flowers. This venerable institution persisted until the French occupation of Belgium in 1794. In 1822, a number of the former members reconstituted the society under the more appro- priate name of the Societé de Flore, and it still exists today as a part of the Societé Royale Linnéenne et de Flore. Under the French occupation the government established a botanical garden in Brussels in 1797, which was located in the grounds of the palace known as the ‘“‘Ancienne Cour.” This garden was intended primarily for the instruction of students in the Ecole Centrale. It had a considerable number of rare plants arranged by the Linnaean system. In 1825, four botanists, headed by J. B. Meeus, approached the burgomaster for permission to found a company to build a new botanical garden. An area of about 15 acres in a good part of Brussels on the newly proposed Rue Royale was at that time the site of a market-vegetable garden. This was bought by Meeus for 43,537 frances. The society was formally approved by a royal decree May 28, 1826, under the name Societé Royale d’ Horti- culture des Pays-Bas. A competition was held for the design of a building, and that by the French painter Geneste was chosen. The building, still standing today, is a fine example of classical architecture. The construction of buildings proceeded rapidly. The green- houses were already finished in 1827, ready to receive the plants from the garden of the Ancienne Cour, which was being abandoned 60 AMERICAN FERN JOURNAL after a disastrous fire in 1826 and the consequent reconstruction of the court. In 1814, near the end of the Napoleonic period, the powers formed the United Netherlands under William I, but this union of the Dutch and Belgians was an uneasy one due to economic problems and the divisive influences of religion and language. The Belgians broke away in 1830 and Dutch troops sent in to quell the uprising lodged in the new greenhouses, causing great damage. There perhaps was no herbarium in the botanical garden, the only herbarium being the Rijksherbarium, at that time in Brussels under the directorship of Blume. When the Dutch withdrew in 1830 they took along the state herbarium to Leiden, where the Rijksherbarium still is, a circumstance that shows how much herbaria may be valued even in times of war and stress. Later, the new king of the Belgians, Leopold I, of faxe-Coburg, gave the garden the collections of herbarium specimens that belonged to the royal court. After the independence of Belgium, the name was officially changed to Societé Royale d’Horticulture de Belgique. The new garden was financed by the selling of shares, for which the stockholders were to receive dividends. The city and govern- ment agreed to a subsidy, but the amount was small, and so the only way that the society could make money was by the sale of plants. Thus almost by necessity the garden was forced to have plants that would grow easily in Brussels and which would sell well, and this consequently led to the neglect of strictly botanical work. Even so, severe financial troubles in 1841 caused a move to dissolve the society, which was in debt for over 203,417 francs. The salvation of the botanical garden was mostly the work of one of the giants of Belgian botany, Barthélmy Charles Joseph Du Mortier. He was born in Tournai April 3, 1797. He did not receive any formal training in botany, but nevertheless devoted himself to it wholeheartedly from about the age of 19, when he began collecting plants in 1816. He married also at the age of 19 and was devoted to his wife, by whom he had eight children, foi the next 60 years. He must have had independent means and some Social status because he worked independently studying ana FERNS IN EUROPEAN HERBARIA 61 writing on botany until 1831, at which time he was elected to the Chambre des Représentants, the equivalent of our House of Representatives. He continued to be reelected for the next 47 years until the time of his death in 1878 at the age of 81. In the early nineteenth century the Linnaean system was still dominant but that of Jussieu was gaining adherents. Du Mortier considered the Linnaean system analytical and that of Jussieu synthetic, and he wished to amalgamate them, an idea that seems rather impossible on the face of it. However, as Crépin remarked, because of his lack of training he did not hesitate to discuss the most difficult problems in botany and zoology. When he was 24, Du Mortier personally published his “Commentationes Botanicae”’ (1822), in which some of his ideas were expressed, a book that is said to have caused a sensation in Belgian botanical circles. The part that is remembered now is the last chapter, in which the old cumbersome genus of hepatics Jun,ermannia was broken up into several, a view that is certainly now considered correct. Du Mortier’s work was original, and he was doubtless much dis- turbed to find out in 1831 that Raddi had anticipated him by publishing a similar segregation of Jun,ermannia in an obscure Italian publication in 1820. Nevertheless, he continued working on hepatics, publishing his “Sylloge Jungermannidearum Europae” in 1831 and more than 40 years later the complete monograph ‘“Hepaticae Jungermannidearum Europae” (1871). Du Mortier worked also on phanerogams, especially Scrophula- riaceae, Batrachium, Salix, and Gramineae. His ‘“Florula Belgica Prodromus’’ (1827) was intended as the forerunner of a complete flora of Belgium, something not in existence then, but he never finished this. Before writing this, Du Mortier visited the Linnaean Herbarium, then in London in the custody of Sir James Edward Smith, to study types, one of the rather few early botanists who availed themselves of this privilege. At the same time he also made an expedition to Scotland to see the elder Hooker. [In 1826, a trip to Scotland probably really deserved the term “expedition.” At the International Botanical Congress in Paris in 1867, Du Mortier presided at the sessions on nomenclature, the 62 AMERICAN FERN JOURNAL principal interest of the Congress and one that took up over half of its time. In 1872 he was made a count by Léopold II. The medal struck for him has below the portrait the words ‘Felix qui potuit rerum cognoscere causas” [Happy is he who can know the eauses of things]. On his 81st birthday in 1878, his statue was unveiled in the botanical garden, a bust that is still there in a place of honor. Du Mortier, as commissioner in charge of scientific establish- ments, was able to persuade the government to double its sub- vention of the botanical garden, provided that the society would renounce its right to dissolution without the consent of the govern- ment. ‘Lhe society was authorized to sell some of its grounds on the side of the present Gare du Nord in order to pay its debts. A period of prosperity then began, especially after H. G. Galeotti was selected as Director in 1853. Henri Guillaume Galeotti wes born September 10, 1814, in Paris, France, of ltalian parents. He was educated as a geolo- gist, and published something on the geology of the region of Brabant at the early age of 20. This brought him to the attention of the Vandermaelen Erothers, the proprietors of a geographical institute and museum in Brussels. They sent him in September, 1835, to Mexico to study the geology and to collect plants, both living and dried, which Galeotti did vigorously, assembling more than 8,000 numbers of herbarium specimens in five years. He began his collecting in the rich region of Mirador and Jalapa in Veracruz. He was perhaps the first botanist to climb the high mountains Orizaba and the Cofre del Perote. In July, 1836, he collected in Real del Monte, Hidalgo, with the German botanist C. Ehrenberg. The next three years were spent on the plateau of Mexico and in the west. In 1839 he went to the southern state of Oaxaca, where he obtained his richest collections. Before returning to Europe in 1840, he established a botanical station in Mirador which was used by later collectors. On his return to Belgium he was made Administrator of the Societé Royale de Flore and later editor of the Journal d’ Horti- culture Pratique. He lived in Louvain, where he became a natu- FERNS IN EUROPEAN HERBARIA 63 ralized Relgian in 1843. His wonderful collections were studied by some of the foremost botanists of the time, the cacti by Lemaire, the grasses by Trinius, and the orchids by Achille Richard. Many of the ferns and phanerogams were published on by the Professor of Botany in Louvain, Martin Martens (1797-1863). Galeotti himself although sometimes a co-author seems not really to have been himself a taxonomist. Unlike most early collectors, Galeotti collected many duplicates and sold them widely, and so they are in most of the large herbaria. Dr. R. Tournay, documentation chief in Brussels, tells me that the Galeotti specimens belonging to Martens were held by the Martens family until 1932, when Pierre Martens, greatgrandson of M. Martens, gave them to Brussels. However, the ‘Index of Collec- tors” of the “Index Herbariorum’” (Regn. Veg. 9, II(2): 214. 1957) indicates that the original and largest set (7,000 specimens) isstill in Louvain, which is a matter that ought to be looked into. Galeotti collected only in Mexico and a few specimens in Cuba but is often cited as the collector of plants from Brazil, Venezuela, and Colom- bia, but these plants were actually collected by Linden, and Gale- otti was merely the distributor of the Linden plants. In 1853, he was appointed Director of the Jardin Botanique de Bruxelles, a post that he retained until his death from tuberculosis in 1858 at the age of 44, a relatively early age considering that as a group Belgian botanists seem to be exceptionally long lived. During his adminis- tration an herbarium was started with Galeotti’s own collections as a basis, to which were added those of Linden, Funck, Schlim, Ghiesbreght, Lindig, and the complete Brazilian herbarium of Peter ( laussen. The botanical library was also enlarged. After the death of Galeotti, A. J. Schram became the Acting Director, and J. E. Bommer was appointed as conservateur (eura- tor). Mr. Schram may not have had the best judgment. The erec- tion of an aquarium was a costly mistake. When Schram left, Bommer was apparently the Acting Director. The necessary re- placement of the heating equipment in 1862 caused some financial embarrassment, so that again the stockholders received no divi- dends. The administration tried without success to get additional 64 AMERICAN FERN JOURNAL subsidies from the city and government. In 1865, the garden had tempting offers of up to 6,000,000 francs for permission to allow roads to run through the garden, and so there was again a deter- mined move to dissolve the society. Again Du Mortier, who was still powerful in the government, came to the rescue. He per- suaded the government to buy the garden for 1,000,000 francs, not a large sum even at that time for such a valuable property. However, it was doubtless large enough for the shareholders to receive their equity. The garden was then renamed the Jardin Botanique de |’ Etat [in Flemish, the Ri jksplantentuin], a title that it retained until 1967, when the name was changed for some unexplained reason to the Jardin Botanique National de Belgique [Nationale Plantentuin van Belgie]. The official establishment of the Jardin Botanique de I’Etat in 1870 was followed a year later by a most propitious occurrence. The famous Bavarian botanist Karl Friedrich Philipp von Martius (1794-1868) had assembled a very large private herbarium, in excess of 113,000 specimens, which was offered by the family to the government of Bavaria for purchase, but the government very short-sightedly refused, because it had already the Brazilian herbarium of Martius and Spix. The family therefore wrote to Anton Spring, the monographer of Lycopodium and a former student of Martius, who formed a committee consisting of him- self, Alphonse De Candolle, and August Eichler. De Candolle wrote a flattering letter to Du Mortier, ending: ‘“‘J’ose insister, parce qu’il est bien rare qu’un Président de Societé botanique et un botaniste consommé soit en méme temps un politique d’un pays. Ce sont des conjonctions aussi rare que celles de certain corps célestin. La Science doit en espérer beaucoup.”’ Du Mortier Was persuasive as usual and the Belgian government purchased the Martius Herbarium in 1870 for 32,000 francs, a ridiculously small sum even for those days for an herbarium containing thou- sands of types and unique specimens. The herbarium was delivered to the state botanical garden in 1871, where it forms the real basis today of the general herbarium. The collections in this herbarium, which were enumerated by Eichler in his paper on the Herbarium FERNS IN EUROPEAN HERBARIA 65 Martii, are far too numerous to list here. They include historic specimens from all parts of the world. The state garden has had very few Directors since its establish- ment a century ago, principally Crépin (1876-1901), Durand (1902- 1912), De Wildeman (1912-1931), Walter Robyns (1931-1966) and Fernand Demaret (1966—present). After Provisional Directors Bommer and Dupont, Francois Crépin was appointed Director in 1876. Crépin was the antithesis of Du Mortier, not a politician and outgoing public figure but a retiring man content with his work with plants. He was born in Rochefort, October 30, 1830, and never received a degree or any formal training in botany. While he was working as a postman at the age of 18 he began botanizing in his spare time, and at 20 quit the post office to devote himself wholly to botany without any sponsor or any hope that he would ever have an official position. For the next ten years he collected and studied the plants of Belgium, and his resulting book, the “Manuel de la Flore de Belgique” published in 1860 immediately established his reputation. Because of its compact size and keys, the work went through several editions and enlargements and was the standard field-guide for two generations of Belgian botanists. His friendship with Louis Van Houtte stood him in good stead. Van Houtte had been employed in the Ministry of Finance. However, when one day he was requested to undertake an im- portant mission over the week-end he replied that it was impossible since he had a date to see the flowering of a rare cactus. The reply did not please the Minister, and from then on Van Houtte was free to devote himself wholly to observing plants, something indeed fortunate for botany since he became one of the foremost protagonists of horticulture and botany in Belgium and indeed in all of Europe. In September of 1861, Michel Joseph Scheidweiler (1799-1861), then in the Institut Horticole of Gentbriigge, died leaving vacant the chair of botany and agronomy that Planchon had for a time occupied. Van Houtte was able to get Crépin this professorship which meant that Crépin immediately had to improvise courses in botany, a task that he did creditably. However, he had little 66 AMERICAN FERN JOURNAL interest in garden plants, and Van Houtte’s continuous impor- tunities to name his greenhouse plants caused a break in their friendship. Therefore, Crépin left Ghent in 1871 to become the curator of the paleontological section of the Musée Royal de V Histoire Naturelle de Bruxelles. The government had purchased a collection of plant fossils from E,. Coemans, and Crépin was delegated to name them. In 1872 he was elected to the Académie des Sciences de Belgique and in 1876 became the Director of the Jardin Botanique de I|’Etat. The staff near the beginning consisted of Bommer, Cogniaux, and Marchal, to which were added sometime later Delogne, T. Durand, and De Wildeman, a distinguished staff indeed. The working room was at the time the large room now containing some offices at the sides and a wood exhibit in the center. Crépin sat in the middle at a big table on a dais where he could survey the work of his staff. On his left at the most primitive kind of tables barricaded by stacks of books and cartons of herbarium specimens sat Bommer, Elie Marchal, and the bryologist Charles Delogne, and on the right Sonnet, Vindevogel, Lubbers (the Head-Gardener), and T. Durand. De Wildeman, as only a volunteer at this time, sat in 1887 at a table a little to the rear. Working conditions must have been difficult, for at first there was no illumination other than the natural light from the windows, and Brussels is not notably bright during the long winters. Only in later years was gaslight installed. Yet the amount of work accomplished was enormous. Crépin himself worked on the genus Rosa, a task on which he spent 40 years, and on which he published over 1,500 pages, but even so his work was never fully completed. His rose herbarium of over 40,000 sheets is kept today as a special unit. Crépin retired because of poor health in 1901 and died in 1903. He was described on a field-trip in 1893 by a journalist as a vigorous sexagenarian, a confirmed bachelor, with a voice sweet and low, who looked like a retired army major except for his modest and reserved attitude, always with his eyes on the ground as if looking at something [which does not seem at all strange to a botanist]. A statue of him was erected in 1904 in his birthplace in Rochefort FERNS IN EuROPEAN HERBARIA 67 in the square named in his honor, the Place Frangois Crépin. For a portrait of Crépin see the biography in Bull. Jard. Bot. de l’ Etat Brux. 1: 281-317. 1905. Fascicle 2 of volume 9 of the same journal is devoted to the celebration of the centenary of Crépin’s birth. Two of the staff members deserve special mention. Célestin Alfred Cogniaux (1841-1916) was born in Robechies. He was appointed Aide-naturaliste in 1872 in the newly organized Jardin Botanique de l’Etat, where he remained for a number of years. He was a prolific worker on three difficult plant families, writing the Cucurbitaceae (1878), Melastomataceae (1883-1888), and Orchidaceae (2037 pp., 1893-1906) for Martius’ “Flora Brasil- iensis,”” and the Cucurbitaceae (602 species, of which 221 were new, 1881) and the Melastomataceae (2,730 species, of which 792 were new, 1891) for DeCandolle’s ‘“Monographiae Phanerogamae.” Finally in his later years, 1909-10, he published an account of the orchids of the West Indies in Urban’s “Symbolae Antillanae.” His valuable herbarium was sold to Brussels. Elie Marchal was born March 1, 1839, at Wasigny, France, of Belgian parents. In 1871, he was chosen by Bommer as Aide- naturaliste in the Jardin Botanique de |’Etat, where he worked on mosses and fungi. He was in charge of the plantings in the garden, where he established sections for ornamental plants, edible plants, medicinal plants, and plants of technological im- portance, all these in addition to the plants of primarily botanical interest. This was in opposition to some prominent Belgian botan- ists like Kickx, Morren, and Martens, who wanted only plants of botanical interest. He was ahead of his time in insisting that all the plants were properly ticketed with their names and acquisition data. In his later years he worked on Araliaceae, and prepared the account of this family for the “Flora Brasiliensis.” For reasons of poor health he retired in 1899, but nevertheless lived up until 1923, when at the age of nearly 84 he was still interested and working on sexuality [of the mosses]. The next Director was Théophile Alexis Durand, one of Belgium’s outstanding botanists, who was born in 1855 in St. Josse ten Noode. He explored the region of Liége and published 68 AMERICAN FERN JOURNAL his first paper on the flora in 1874 at the age of 19. At the sug- gestion of Elie Marchal he went to Switzerland in 1877 to study the alpine plants, and published a series of papers (1881-1887) with his friend Henri Pittier, at that time Professor of Botany at Chateau d’Oex. In 1879 he started work as a volunteer in the Jardin Botanique de |’Etat. By his continuous work and the care he gave to all the tasks confided to him he won the regard of Crépin and was appointed to the staff as Aide-naturaliste in 1881, and later became the curator. At first he worked on the bryophytes of Belgium with Delogne, but soon became interested in com- piling an index of genera, the result being the “Index Generum Phanerogamorum,” a work that made him famous and one that is still most useful to the herbarium botanist. He then col- laborated with B. D. Jackson in preparing the first supplement to the “Index Kewensis.” In the meantime he continued floristic work with Pittier on the “Primitiae Florae Costaricensis.” In 1895, at the request of the government, it was decided that the garden should undertake the study of the flora of the Belgian Congo, and Crépin delegated Durand and De Wildeman to the task. This collaboration resulted in the formation of the “Annales de Musée du Congo” and the publication of many important papers on the plants of that region until eye trouble forced his retirement in 1908. He died in 1912.1 The next Director, one of the brightest lights of Belgian botany, was Emile De Wildeman, who was born in Brussels, October 19, 1866, where he died July 24, 1947. He became a volunteer worker in the botanical garden in 1887 and was taken officially on the staff in 1890, where he remained for 41 more years of service before retiring in 1931. At the time he was one of the first students of the famous professor Léo Errera (the formulator of what is called “Errera’s Law” that a cell membrane tends to assume the same form as a weightless fluid layer would assume under the same conditions), who had just started a botany department in the 1 4 - ’ 42): ovis 188. Ts Dune ee Bull, Jad, Bot. de VEtat Brux American Elias M ; eR ae ¢ as M. Durand, whose historic herbarium containing Rafinesque is now in Paris. FERNS IN EuROPEAN HERBARIA 69 University of Brussels in 1891. De Wildeman received his doctorate in 1892 with a thesis on cell division. In his early years he worked and published on algae, but in 1895 Crépin decided to charge him with the preparation of a flora of the Belgian Congo, a task on which he spent the rest of his life. The herbarium collections that had been accumulated by the Etat Independant du Congo were turned over to him, the beginning of the present tremendous collection of Congo plants numbering more than 500,000 sheets in 1948. De Wildeman became the foremost authority on the flora of the Congo, and was very likely the last botanist to describe thousands of new species [because there are no more completely unknown parts of the globe]. It was perhaps not necessary to describe quite so many, because recent studies have shown that De Wildeman did not always take into account the species that had been described from other parts of tropical Africa, and also had perhaps too narrow a species concept, but even so most of his species are good ones. He was one of the most prolific workers ever, with a bibliography of more than 1,400 titles, ranging from brief notes to large books. His final work, entitled rather sadly, “Stérilité ou Viellissement et Disparition des Espéces Végetales”’ was published in two volumes a year after his death at the age of 80 in 1947. He was the Secretary-General of the International Botanical Congress that was held in Brussels in 1910. In 1912 he became Director of the Jardin Botanique de |’Etat.” Young botanists and perhaps especially the old too) should ponder his words: “Ne cherchons pas & faire oeuvre définitive! Ne nous ber- ¢ons pas du fallaceiux espoir de produire des travaux sans dé- fauts. La perfection n’est pas de nétre monde?” During the regime of W. Robyns, the next Director, it became apparent that the old building and greenhouse were inadequate and that the garden was suffering from the city air. It was there- fore arranged in 1938 that the garden be moved to the Domaine de Bouchout, in the country near Meise, a few kilometers north of Brussels. This is a large estate of over 200 acres, with a lake and an imposing chateau, formerly a royal residence. The advent of ? For a biography, see Bull. Jard. Bot. de l’Etat 19: 1-31. 1948. 70 AMERICAN FERN JOURNAL the war in 1939 naturally stopped any move to the new quarters. About 10 years later work was started, and now the huge green- houses are finished and all the plants moved from the old ones. A magnificient new administration and herbarium building is awaiting occupancy. The new garden is not so accessible as the old one in the center of Brussels, but is well worth a visit. In addition to the greenhouses the grounds themselves contain many fine old trees dating back a century or more. Certain areas are being left in a natural state for the wild vegetation. The new garden may be reached rather easily by taxi or bus. There are however, I believe, no accomodations nearby where visitors can stay for longer periods. Robyns continued with De Wildeman’s work on the Congo, publishing volumes on the grasses and a “Flore des Spermato- phytes du Pare National Albert” in three volumes (1948-1955). He also started the publication of the “Flore de Congo Belge et du Ruanda-Urundi,” of which ten volumes appeared between 1948 and 1963. This work is being continued, but now in the form of separate fascicles for the different families, which can thus be issued as they are ready without any set taxonomic sequence. The present Director, Fernand Demaret, an authority on the bryophytes of the Congo, first came to the garden in 1936 as an assistant and became director in 1966. September 1970 being the centenary of the establishment of the garden as a government unit, the garden held a formal celebration of the event. Botanical institutions throughout the world were invited to send representatives for the ceremonies. Since I had been in Brussels in June of that year, I did not feel that I could attend, but I understand that the event was highly successful in every way. FERNS IN BRUSSELS The principal pteridologist in the early years was Jean Edouard Bommer, who was born November 17, 1829, in Brussels. He had no formal education in botany, but was a hard and dedicated worker, at first employed in the Etablissement Vandermaelen, FERNS IN EUROPEAN HERBARIA 71 but by 1856 he was well enough known that he joined the staff of the botanical garden in Brussels as Aide-naturaliste, and after Galeotti’s death he was the curator and later Provisional Director. Although Bommer worked on various groups of plants, he was mostly interested in ferns. His first paper was on the hairs of ferns, a subject that has been somewhat neglected but now realized to be of fundamental importance (Bull. Soc. Roy. Bot. Belg. vol. 1). His most important paper is the “Monographie de la Classe des Fougéres,”’ which is not a monograph but a paper on fern classification that is little known today. However, Bommer’s classification was essentially that adopted by Eichler and later by Engler, and is thus quite similar to the classification adopted still later by Christensen, except for the inclusion of the Cyatheaceae in the Polypodiaceae. He collaborated with the Swiss botanist H. Pittier in publishing on the numerous then recent collections in Costa Rica of Pittier, Tonduz, Biolley, and others, in Durand and Pittier’s ““Primitiae Florae Costaricensis.’”’ At the time of his death February 19, 1895, he was working on a monograph of the genus Adiantum. According to Dr. Lawalrée, the manuscript has disappeared. Strangely enough, however, a synopsis of the pro- posed classification, in Bommer’s own hand, is in the Smithsonian Institution, a Xerox copy of which has now been deposited in Brussels. Bommer can not be accounted an outstanding pteridol- ogist, but his influence is clearly evident in the Brussels herbarium, where his annotations are on many sheets, and where many of the older specimens in the general herbarium are still arranged as he left them.* Among the valuable parts of the old fern collections is the fern herbarium of Peter Claussen, but it seems that little is known of Claussen or his collections. It may be surmised that Linden and Claussen knew each other, since they were in Brazil at the same time. Galeotti must have been close to Linden, and through him with Funck and Schlim, Triana, and Lindig, for all these collections were turned over to Galeotti for the garden and for distribution. Thus Brussels is very rich in these old collections, especially those * For a biography, see Léo Errera, Bull. Soc. Roy. Bot. Belg. 34: 7-21. 1895. 72 AMERICAN FERN JOURNAI from Venezuela and Colombia that formed the basis of G. Metten- ius’ treatment of the ferns in Triana and Planchon’s ‘‘Prodromus Florae Novo-Granatensis” (Ann. Sci. Nat. [Paris] V, 2: 192-271. 1864). The specimens in Brussels with the names in the hand of Mettenius should be considered the holotypes. These collections are partly represented by duplicates or fragments in the Met- tenius Herbarium in Berlin. The Linden ferns are of particular importance. Jean Jules Linden was born in Luxemburg in 1817. At the age of 18 he was authorized by the Belgian government to collect plants in tropical America with Nicholas Funck and Auguste Ghiesbreght as companions. They first went to southern Brazil. On the second trip Linden went alone to Cuba in 1837, and in 1838 to Mexico and Guatemala, where he obtained sizeable collections, especially in Tabasco and Chiapas. His third and most profitable trip was made with Louis Joseph Schlim in 1841 to Venezuela and Colom- bia, where he remained until 1844. In 1844-1845, Linden collected extensively in Cuba, especially in the eastern province of Oriente. On his return he established a nursery in Luxemburg in 1845, which was moved in 1851 to Brusseis, where it became one of the foremost in Europe. In 1852 or 1853 he was appointed Director of the Jardin Royal de Zoologie et d’Horticulture de Bruxelles. Linden listed his offerings in catalogues that are rare and much prized today because they contain the original descriptions of some of the plants that Linden introduced to cultivation. In 1870 he expanded his establishment by buying the large nursery of Ambroise Verschaffelt in Ghent. Linden’s collections were never completely worked up, although some groups were studied, e.g. the “Orchidaceae Lindenianae” by J. Lindley. Linden sold his plants widely. There is a large set in Brussels, and an unexpectedly large set in the Webb Herbarium in Florence. Linden remained active until the age of 80, when he died in 1898, at the time the Consul-General in Belgium for Colombia. There is an account of his itineraries in Annales Soc. Roy. Agr. Bot. Gard. 2: 205-220, 259-272. 1846. Linden’s associate, Nicholas Funck (1816-1896), was also born FERNS IN EuROPEAN HERBARIA 73 in Luxemburg. He was sent to Mexico with Linden in 1835 to collect, and in 1845 he went with Schlim to Venezuela and Colom- bia, where he assembled large and valuable collections. Un- fortunately many of these were lost in a shipwreck, but even so there are some 4,650 numbers of his collecting in Brussels. A collection of ferns of great importance is the Indian and Malaysian collection of William Roxburgh. I expect to publish a paper on the Roxburgh ferns, and so I shall not comment on them here other than to say that the Brussels set is the most authentic, with the names in Roxburgh’s own hand. The Costa Rican ferns of Pittier, Tonduz, Biolley, Wercklé, and others were worked up by Bommer, and later by the Swiss botanist Hermann Christ. Many of the Christ types from Costa Rica are in Brussels rather than in the Christ Herbarium, now in Paris. It is evident that when J. G. Baker was working on his account of the Cyatheaceae and Polypodiaceae for Martius ‘Flora Brasil- iensis” (vol. 1, no. 2. 1870), that he had on loan the Brazilian specimens from Martius’ herbarium, for they are annotated by Baker and correspond with the names as used in the Flora. In the case of the new species therefore Baker’s types are in the Martius Herbarium in Brussels and not at Kew, where Baker’s other types mostly are. Sometimes Baker removed fragments and these are at Kew. The types of Johann Wilhelm Sturm (1808-1865), who wrote the accounts of the Hymnophyllaceae, Gleicheniaceae, Marattiaceae, Ophioglossaceae, Osmundaceae, and Schizaeaceae for the “Flora Brasiliensis,” are also in the Martius Herbarium in Brussels. The holotypes of the species described in 1824 by H. A. Schrader from the Brazilian collections of the Prince von Neuwied are also in the Herbarium Martius lent ie Brazilian collections to Gustav Kunze, the noted pteridologist of Leipzig, who published his paper “Filices in Martii Herbarium Florae Brasiliensis” in 1839. These specimens were returned to Martius, and thus the holotypes of these Kunze species are still in existence, with the names in Kunze’s hand, fortunately, since Kunze’s own herbarium was destroyed in Leip- 74 AMERICAN FERN JOURNAL zig during World War II. Many of the South African species de- scribed by Kunze have also authentic material named by Kunze in Brussels. The other fern collections include those of Pohl, Karwinski, Blanchet, Luschnath, Glaziou (many numbers), Husnot, Gay, Bertero, and others far too numerous to mention. Naturally, Brussels is the richest herbarium for Belgian ferns, and those of France are well represented, due especially to the fine herbarium of J. Callé and his collaborator E. Walter. Brussels is now fortunate in having pteridologists again for the first time since the late nineteenth century. The Director, F. Demaret, is interested in the ferns of the Congo. Dr. A. Taton also has studied African ferns and has published an account of the Congo Hymenophyllaceae. Dr. André Lawalrée, the present pteridologist, prepared a perceptive account of the ferns for the “Flore Générale de Belgique” (1950) and is now at work on the ferns of the Congo, of which six parts have been published in the “Flore du Congo, du Rwanda et du Burundi,” the Equisetaceae, Psilotaceae, Actiniopteridaceae, and Parkeriaceae in 1969, the Schizaeaceae in 1970, and the Blechnaceae in 1971. Collaborators preparing manuscripts include Kramer (Lindsaeaceae), Launert (Marsileaceae), Bizarri (Selaginellaceae), and Pichi-Sermolli. I have visited Brussels three times, the first in 1954 to study and photograph the types of Martens and Galeotti. Prior to 1842, when their “Mémoire sur les Fougéres du Mexique” was published by the Académie Royale de Bruxelles, very few ferns of Mexico were known. Martens and Galeotti described many new species, to be wrongly identified and actually undescribed at the time. However, sometimes the Martens and Galeotti species belong to very critical groups and they can be known definitely only by the study of the original specimens. Many of the new species have proved to be common, but others have perhaps never been found again. There are many duplicates of the Galeotti collections in Brussels, but it is usually possible to tell which one is the holotype. FERNS IN EUROPEAN HERBARIA 75 In my visits to Brussels I have always been most hospitably received, in 1954 by Dr. Robyns and Dr. Lawalrée, and in 1969 and 1970 by Dr. Demaret and again Dr. Lawalrée. Unfortunately, the ferns along with the general herbarium, including the Martius Herbarium, are now kept in a basement that is not well illuminated and possibly at times too damp. It is to be hoped that they can be moved soon to the new herbarium building that is ready for them. Much of my information comes from the paper by J. E. Bommer “Notice sur le Jardin Botanique de Bruxelles,” which was written at the time when the present garden was first formed (Bull. Soc. Roy. Bot. Belg. 9: 418-455. 1871). Some of the recent information comes from the publication that was issued last year for the centenary “Le Jardin Botanique National de Belgique, 1870- 1970,” which is unsigned, but is by Dr. Demaret, Dr. Lawalrée, and their associates. (To be continued) An Appendageless Psilotum. Introduction to Aerial Shoot Morphology ALBERT 8S. Rourra! Over forty years ago Okabe (1929) wrote that ... “ Psilotum nudum, which grows wild in southern Japan, was once much cultivated in Japanese gardens as an ornamental plant. Slightly over 100 garden varieties were given fantastic names. Today, however, there are unfortunately only a few of them found oc- casionally in greenhouses of botanical gardens or in private gardens.” Since the time that Okabe wrote this apparent epitaph there have been several attempts by Psilotum growing associations in Japan to revive the almost lost art and to gather information on the remaining varieties available. According to a special iversi inoi i Circle 1 Supported by a grant from the University of Illinois, Chicago Research Board. AMERICAN FERN JOURNAL VouuME 61, PLATE 14 i ; 4 a Fig. 1. A DISTAL PORTION OF AN AERIAL SHOOT WITH FIVE BRANCHES OF VARYING LENGTHS BEARING TERMINAL SYNANGIA, THE LATTER WITH MORE T THE TYPICAL THREE LOCULES, X 3.0. Fic. 2. A PSEUDODICHOTOMY, BOTH BRANCHES TIPPED WITH MULTIPLE SYNANGIA, THE LEFT BEARING TWO VISIBLE STUB-BRANCHES, XX 3.0. APPENDAGELESS PsILOTUM te issue in December 1967 of “Horticultural Magazine” (Japan) en- titled ‘Classical Horticultural Plants” there are about 50 clearly identified old varieties of Psilotum plus about 10 unidentified and newly introduced varieties in existence. Among the older clones which may have been grown for more than 200 years there is one known as sei ryu kaku, a descriptive appellation that may be broadly interpreted in the traditional sense as “Green Dragon’s Horns” or in a more modern sense as “Green Giraffe’s Horns” (Rouffa, 1968). With all apologies to Okabe, the name, far from fantastic in this instance, appears rather appropriate (see Figs. 1-8). It is interesting to note that this Psilotum does not produce either sterile or fertile appendages and develops meiotic synangia at the tips of the ultimate branches. For this reason, this Psilotum merits an introduction and further study. The clone was introduced into the United States in 1956 by Dr. John Creech (1957), of the New Crops Research Branch, U. S. Department of Agriculture, Beltsville, Maryland as P.I. 235301 and, along with a number of other Japanese Psilotums, deposited and grown for a number of years at Longwood Gardens. Subsequently, some of the collection (not including this clone) was sent to the New York Botanical Garden. I am particularly indebted to Dr. Donald Huttleston of Longwood Gardens for the special propagation and donation of their Psilotums to our department in November 1967. Up to that time the unusual characteristics of sei ryu kaku were quite unknown to any of us until the plants were unpacked in our laboratory. I am also especially grateful to Mr. Kaname Kato, an eminent horticulturist in Tokyo, for providing me with valuable informa- tion regarding this particular clone and the history and culture of Japanese Psilotums in general. MATERIALS AND METHODS Severed shoot tips were treated in a number of ways for observa- tion with the Cambridge Mark II stereoscan SEM (see figure legends) and included preservation in both glutaraldehyde and AMERICAN FERN JOURNAL VoLuME 61, Pate 15 Fig. 3. A DISTAL PORTION OF AN AERIAL SHOOT WITH A SERIES OF BRANCHES OF VARIOUS LENGTHS: Two LENGTHS OF STUB BRANCHES AND A TERMINAL PSEUDODICHOTOMY WITH SHORT BRANCHES TERMINATING IN SYNANGIA, X 2.0. Fic. 4. AN OVERALL VIEW OF A GROUP OF SYNANGIA-BEARING AND STERILE AERIAL SHOOTS, X 0.8. APPENDAGELEss PSILOTUM 79 in FAA. A study is continuing to determine the best treatment for various conditions. Fresh tips could be observed for periods up to thirty minutes, when vacuum-induced shrinkage became apparent. Psilotums have been grown in our laboratories for a number of years indoors under fluorescent lights. They appear to grow best and produce more sporangia when cool-white lamps are supple- mented with wide-spectrum Gro-Lux lamps in a ratio of 2:1. Light intensities range from 1,000-2,500 ft-c. Sei ryu kaku has responded well to this lighting and grows as vigorously as many of the typical Psilotums. Despite the nearly constant cultural conditions, all Psilotums tend to alternate periods of rapid growth with periods of quiescence. OBSERVATIONS When aerial shoots of sei ryu kaku emerge from the soil they appear relatively smooth and terete. Their often apiculate apex is unobscured by leaf primordia. Therefore one can easily observe the subdividing of the apex into multiple apices (Bierhorst, 1954) often when the shoot is not more than 2 cm in length above the soil (Figs. 10, 11, 13). After its formation, each apex eventually develops its own subtending axis before the final elongation of the total aerial shoot takes place (Fig. 9, 11). The shoot apices continue to subdivide following elongation of the total shoot (Figs. 13, 14). The early subdividing of the emerging aerial shoot apex is synchronous and symmetrical (Fig. 13), but the final sub- divisions of the apices appear to be less synchronized and some may be quite asymmetrical (Figs. 14, 15). This asymmetry is perpetuated in the growth of some of the distal branches. As a result, distal shoots may develop monopodially and produce lateral branches ranging in size from mere ridges, to stub-branches 1 mm in length, to nearly equivalent pseudodichotomies (Figs. 1-5). Equal pseudodichotomies may also develop (Fig. 9). Unequal branches are often, but not always, the hallmark of shoots forming synangia (Figs. 1-8). AMERICAN FERN JOURNAL VouumE 61, PLATE 16 Fic. 5. SHOOT TERMINATING IN STERILE FORKED-TIPPED BRANCH AND A SYNANGIAL BRANCH; GLISTENING DOTS ON SURFACE ARE STOMATE SITES, X 1. Fies. 6-8. SHOOTS TERMINATING IN SHORT SYNANGIA-BEARING PSEUDODICHO- TOMIES. Fics. 7 & 8. SAME SHOOT TAKEN ON OPPOSITE SIDES. Fic. 6, X 3; Fias. 7, 8, X 1.5. Fie. 9. ACTIVELY ELONGATING VEGETATIVE UPPER SHOOTS; NOTE THE ABSENCE OF APPENDAGES, X I. Fic. 10. BASAL PART OF SHOOTS NEAR GROUND LEVEL; NOTE ABSENCE OF STERILE-APPENDAGES AND THE NAKED APPENDAGELESS PSILOTUM Sl The stub-branches are suggestive of the sterile appendages of the typical leafy form of Psilotum. On the whole, shoot vascu- larization of sei ryu kaku appears similar to that of the typical form. Freehand sections taken at suitable intervals along the shoot and stained with HCl-Phloroglucinol reveal that the stub- shoots have a lignified strand branching from the stele extending at least to their base. Those stub-shoots bearing synangia are supplied with a lignified strand to the synangium. In certain respects, particularly in their vascularization, the sterile stub- shoots resemble the sterile appendages of Psilotum complanatum (Stiles, 1910). However, the stub-shoots are not necessarily pro- duced in helical order, although occasional distal shoots may show this arrangement (Fiy. 3). Synangia typically develop at the tip of certain distal aerial branches. Such branches may be of varying lengths (Figs. 1-8). Meiosis has been observed in the synangia, and apparently normal kidney-shaped spores are eventually produced in the locules. The number of locules may be 4 or more rather than the usual 3 (Figs. 1-3, 8). As in typical Psilotums, not all aerial shoots bear sporangia, and there is a tendency for sporangium-bearing shoots to occur in groups (Fig. 4). Spore production (on a per-aerial shoot basis) may be lower for sei ryu kaku than for the typical Psilotum SHOOT APICES, X 1.2. Fic. 11. CLosEUP OF BASAL PORTION OF SHOOT ABOUT 4 MM FROM SOIL LEVEL AFTER FORMATION OF MULTIPLE APICES, THE UPPER SHOOT SYSTEM BEGINNING TO ELONGATE, X 2.5. Fic. 12. ME1oTIC METAPHASE SHOWING 50 BIVALENTS. SEM PHOTOGRAPHS OF SHOOT APICES: FIG. 13. Group OF MULTIPLE SHOOT APICES AT SAME LEVEL AS Fig. 11; Frxep in FAA; 50% GLYCERINE 24 HRS.; GOLD COATED; NOTE THE EARLY SUBDIVISIONS GENERALLY EQUAL AND SYMMETRICAL, BUT LATER AND INCIPIENT ONES LESS SO, x 14. Fie. 14. Group OF SUBDIVIDING SHOOT APICES FROM A SLIGHTLY LATER STAGE THAN QUITE UNSYMMETRICAL; FRESH AND UNCOATED, X 23. Fie. 15. Grou ELONGATING SHOOTS AT A SOMEWHAT LATER STAGE OF GROWTH THAN Fie. 13, THESE APICES PERHAPS REPRESENTING THE LAST SUBDIVISIONS PRIOR TO THE COMPLETE AND FINAL ELONGATION OF THE SHOOT; NOTE INITIAL SYMMETRY AND LATER ASYMMETRY; CF. Fic. 5; rrxep IN FAA; 50% GLYCERINE 24 HRS.; GOLD COATED, X 27.5. 82 AMERICAN FERN JOURNAL where synangia are produced by a series of lateral fertile-append- ages along the distal ends of the aerial shoots. There is evidence of meiotic irregularities. Whether those observed were induced by environmental conditions under which the plants were grown (Okabe, 1929; Barber, 1957), or for other reasons, is not known. In initial chromosome counts of SMC metaphases pretreated with oxyquinoline and stained with aceto- carmine, meiotic bivalents ranged from about 46—56 (e.g. Fig. 12). Sei ryu kaku is therefore a diploid. Discussion The entire scientific literature of Psilotum nudum and P. complanatum that deals with the origin and relationships of this genus has never considered any type of aerial shoot other than the one that produces reduced leaflike appendages of two inter- grading types: (i) a sterile series of awl-shaped appendages on the lower extremities of the shoot, and (ii) a fertile series of forked appendages on the upper extremities of the shoot which is associ- ated with the development of synangia at each appendage (Bier- horst, 1956; Roth, 1963a, b). In sez ryu kaku, however, the lower extremities do not have leaflike appendages (Fig. 10). When the upper extremities of the shoot are differentiated they are also appendageless (Fiy. 9), but stub-branches may be produced (Figs. 2, 3) by asymmetrical growth. These short stub-branches may resemble the sterile appendages of typical Psilotwm, but are not produced in a continuous series nor is their number or size consistent. In this variety the synangia are borne only on the tips of branches (F igs. 1-8). The genus Psilotum and its companion genus T'mesipteris have long been considered modified subtropical and_ tropical relics of the early land plants of the lower Devonian. This was due m part to the apparent nature of their sporophytes, which seemed to bear little resemblance in growth and structural patterns to those of other primitive plants. Recently, however, Bierhorst (1968a, b, 1969) has taken another look at the problem and has shown a number of structural and developmental similarities APPENDAGELESS PsILOTUM 83 between T'mesipteris especially and the enigmatic New Caledonian endemic fern Stromatopteris moniliformis. Noting greater simi- larities between the two genera of the Psilotaceae and Strematop- teris than the latter possessed in regard to the family Gleichenia- ceae, to which Stromatopteris had originally been taxonomically assigned, Bierhorst (1968a) proposed that this apparently anomalous fern be placed in a family of its own, the Stromatop- teridaceae. He further stated that the Psilotaceae and the Stromatopteridaceae were too closely related to be separated into separate orders. As a result he transferred Psilotum and Tmestpteris (Psilotaceae) into the order of ferns (Filicales). As a corollary, the aerial axes of Tmesipleris and Psilotum would then be considered homologous with the aerial ‘frond’ of Stromatopteris. Bierhorst (1969) also postulated that the radially organized aerial axes of Psilotum, rather than being primitive, were actually specialized and derived from the winged, leaflike condition of Tmesipteris. These sweeping conclusions were based on an impressive amount of morphological and anatomical evidence, much of it gleaned from Bierhorst’s own indefatigable collections in the field. The phytochemistry of the Psilotaceae has been recently enriched by the discovery of a phenolic glycoside, psilotin, that was isolated and identified from Psilotum nudum (McInnes, Yoshida & Towers, 1965) and T'mesipteris tannensis (Tse & Towers, 1967). According to Tse & Towers the compound has not been identified in representatives of the Lycopodiophyta. There is a possibility that the phylogeny of the Psilotaceae, long dormant as a subject for scientific inquiry, may well again be- come a subject of lively controversy. It may be that, in light of Bierhorst’s conception that Psilotum has a derived radial shoot or axis, the unusual characteristics of set ryu kaku may or may not have a direct bearing on the determi- nation or confirmation of the phyletic affinities of Psilotum. This clone is of more immediate morphogenetic interest because it performs routinely the production of terminal synangia, a phenom- enon that occurs only quite rarely, or under environmental stress, in most Psilotum genotypes. In typical Psilotum, spontaneous 84 AMERICAN FERN JOURNAL proliferation of the fertile appendage into a short branchlike struc- ture was reported by Sykes (1908) and Bierhorst (1952). Typical Psilotum grown under normal or greenhouse conditions have produced synangia in apparently terminal positions (Iabbri, 1963; Abraham & Ninan, 1965; Rouffa, unpublished observations). Also, Bierhorst (personal communication) has noted the existence of a native New Zealand clone in which the synangia are borne at the tips of distinct branches. Such occurrences could certainly strengthen the theory that the forked fertile appendage was evolved from a reduced branch or axial system with terminally produced sporangia. (Bierhorst, 1956; Rouffa, 1967). Indeed the resemblance of the elongated synangium tipped shoots of sei ryu kaku to the aberrant, elongated fertile appendages tipped with synangia produced for at least six years in the author’s laboratory on a typical variety of Psilotum is rather striking (see Rouffa, 1967, fig. 4). The latter aberrant fertile appendages were induced under conditions of low light intensities, red spectrum deficiency, and high temperatures. The effect, although pro- duced quite regularly on an annual basis, appeared unstable since there was a tendency for the same shoot to return to the formation of the normal fertile appendage-synangium structures, even though the inducing environment remained unchanged. To explain this, I postulated that typical Pszlotum still carries in its genome an ancestral set of genes responsible for the timing sequence of meiosporangium development at the tips of certain branches, but that some form of gene repression (possibly genes acquired through polyploidy) “modernizes’”’ the expression of the branch-sporangium combination to that of a fertile appendage- sporangium combination. I further suggested that, somehow, repressing genes themselves were themselves repressed tempor- arily at the time of the experiment by the extreme stress of an unusual environment, thereby unmasking temporarily the an- cestral condition where sporangia are produced terminally on elongated shoots. Set ryu kaku, however, normally produces synangia on the ultimate tips of shoots (both long and short) and does not produce APPENDAGELESS PsILOTUM 85 appendages. According to the above hypothesis, the “modern- izing” genes are either continually repressed, or they are not present in the genome in this clone. Future studies on chromosome irregularities in this clone might shed some additional light on these possibilities. It cannot be denied that this much prized Japanese exhibition Psilotum looks for all the world like something out of the Devonian. However, in the face of new evidence, we must exercise caution in assuming that this variety, or any other form of Psilotum, is a direct or relic descendent from that period. Nevertheless, there is a distinct possibility that in sei ryu kaku some genetic change, as yet unilluminated, has stripped away several hundreds of millions of years of evolutionary modification and there is a re- mote chance that it represents one of the last of a nearly extinct race of Psilotum. Finally, it may be that we now have been provided the oppor- tunity to learn from its relatively simple structure how the early land plants solved their developmental problems. We are now gathering for this purpose an array of variant genotypes of Psilotum which should prove most useful for genetic and morphogenetic analyses and for phytochemical studies. LITERATURE CITED ABRAHAM, A. and C. A. Ninan. 1965. Morphological and cytological notes on Psilotum nudum (L.) Beauv. Caryologia: 7-539. Barger, H. N. 1957. Polyploidy in the Psilotales. Proc. Linn. Soc. New uth Wales 82: 201-207. Brernorst, D. W. 1952. The Sera et anatomy es hes phylogeny of the Psilotaceae. Ph.D. Thesis, University of Minn —————. 1954. The origin of branching in the ice ag of Psilotum nudum. Virginia J. Sci. 5: 72-78. ————. 1956. Observations on the aerial appendages in the Psilotaceae. Phytomorph. 6: 176-184. oe + ee On the eae (Fam. Noy.) and on the Pa tarene Phytomorph. 18: 232-268. i ree bhi on Schizaea and Actinostachys spp., chain A. oligostachys, sp. nov. Amer. J. Bot. 55: 87-108. S6 AMERICAN FERN JOURNAL Birrnorst, D. W. 1969. On Stromatopteris and its ill defined organs. Amer. J. Bot. 56: 160-174. Creecn, J. L. 1957. Plant explorations. Ornamentals in southern Japan. U.S. partment of Agriculture, Agr. Res. Serv. 34-1: 1-55. Fassri, F. 1963. Terminal sporangia at the apices of Sooty in two plants of — nudum (L.) Beauv. A preliminary report. Giorn. Bot. Ital. 70: 591-595. aaa G, x ee and G. H. N. Towers. 1965. A phenolic gycoside m Psilotum nudum (L.) Griseb. Tetrahedron 21: 2939-2946. NINAN, c. A. 1956. Cytology of Psilotum nudum (L.) Beauv. (P. triquetrum w.). La Cellule: 57: 307-318. OKABE, : 1929. Uber eine tetraploide Gartenrasse von Psilotum nudum, Palisot de Beauvois (= P. triquetrum, Sw.) und die tripolige Kern- ea Bi In ihren Sporenmutterzellen. Science Reports, Tohoku Im- rial University, IV, 4: 373-379. Rovurra, oe S. 1967. Induced Psilotum fertile-appendage aberrations. Mor- phogenetic and evolutionary implications. Can. J. Bot. 45: 855-861. —————. 1968. An appendageless form of Psilotum. Amer. J. Bot. 55: 714. (Abstr.) Rorn, I. 1963a. Histogenese der Luftsprosse und Bildung der “‘dichotomen”’ Verzweigungen von Psilotum nudum. Adv. Front. Pl. Sci. 7: 157-179. 963b. Histogenese und Tale nag tes ee ee Blatter von bile tum nudum. Flora (Allgem. Bot. Zei 90-11 Stites W. 1910. The structure of the aerial en a eae flaccidum, Wall. Ann. Bot. 24 (O.S.) : 373-387. Sykes, W. G. 1908. A note on an mae found in Psilotum triquetrum. nn. Bot. 22 (0. S.) : 525-526. Ts, ae pare G. H. N. Towers. 1967. The occurrence of psilotin in Tmesip- teris. Phytochemistry 6: 149, DEPARTMENT oF BIor ScIENCEs, UNIVERSITY OF ILLINOIS AT Sareaap Circe, Box 4348, Cuicaco, ILLINoIs 60680. VASCULARIZATION OF FERN LEAVES S7 Vascularization of Fern Leaves Ropert C. LomMAsson AND C. H. Youna, Jr. The subject of fern leaf venation is a very old topic and it has been dealt with by many authors during the nineteenth and twentieth centuries. Most of the discussions of fern leaf venation concentrate on its qualitative aspects resulting in descriptions of various types of leaf venation. Considerably less attention has been devoted to the quantitative aspects of fern leaf vascular- ization. This report will give a brief summary of some of the measurable aspects of the vascular system within the leaves of certain rather common ferns. The collections of the ferns used in this study came from two sources. Those collected out-of-doors came from Giant City State Park or from Jackson Hollow, six miles northeast of Simpson, Illinois. Material was taken with the permission and under the supervision of Dr. Robert H. Mohlenbrock of Southern Illinois University. Other ferns were obtained from the Department of Botany greenhouse at the University of Nebraska. The fern leaves were studied from slides made of cross sections of the material which had been stained with Delafield’s hematoxy- lin and safranin. Whole mounts of cleared leaves or leaflets or parts of these were also studied to obtain vein patterns and angle of branching. All measurements recorded in this study were made by means of an ocular micrometer. All those listed in Table I are averages of many separate measurements. Leaf thickness was measured at veins rather than between veins. Intervascular interval was measured from the vascular tissue of one bundle to the vascular tissue of the next adjacent bundle; thus it includes any specialized parenchyma or bundle sheath cells. Vein width includes only the width of the conductive tissues of a vascular bundle. Angle of branching refers to the branching of vascular bundles from the midvein of the leaf or segment and not to the ultimate branchings in the lamina. It was measured in linear units and the angle SS AMERICAN FERN JOURNAL computed as a trigonometric function. Dissection of leaves as used in this study was an arbitrary rating scale which divided Tass [. Summary or Lear MEASUREMENTS OF SOME COMMON FERNS LT IVI VW ABin Species Source inp inn inp degrees D Botrychium virginianum I 260 747 85 39 5 Osmunda cinnamomea I 125 426 32 41 3 Trichomanes boschianum I 98 799 45 51 i Pteris vittata G 108 463 53 43 2 Pteris cretica G 174 791 79 39 2 Cheilanthes lanosa I 135 281 28 43 4 Pellaea atropurpurea G 195 96 45 41 4 Adiantum pedatum G 56 622 38 48 5 Adiantum capillus-veneris G 42 323 20 32 5 Nephrolepis exaltata G 176 561 45 38 2 Woodsia obtusa AE 177 429 39 37 3 Polystichum Sean I 286 496 44 53 2 yrtomium falcatu G 336 744 78 42 2 Dryopteris den a G 108 404 43 42 3 Dryopteris marginali: is I 171 607 42 42 a yo losa I 110 633 38 40 5 Dryopteris hexagonoptera I 99 581 18 45 3 Athyrium filiz-femina I 138 437 30 33 4 Asplenium trichomanes I 429 30 34 2 Asplenium platyneu I 140 500 38 35 2 amptosorus rhizophyllus I 219 984. 57 62 0 Polypodium vulgare G 176 736 78 43 1 olypodium virginianum I 200 681 42 44 1 Phlebodium aureum G HA 1208 69 44 1 I = Southern Illinois; G = Greenhouse, ae of Nebr.; LT = leaf thickness; IVI = Intervascular interval; VW = vein width; AB = angle of a ige D = Dissection of leaf: 0 = entire; 1 = hasnt 2 = pinnate; 3 = nate-pinnatifid; 4 = twice-pinnate; 5 = more than twice-pinnate. fern leaves into six groups. These were designated by a numerical seale from 0, which represented entire leaves, to 5, which included those which were more than twice pinnate. VASCULARIZATION OF FERN LEAVES 89 Lear THICKNEss In general, the leaf thickness at vascular bundles is greater than it is between them. In five species the leaf thickness at the vascular bundle exceeded the sum of leaf thickness between veins and the vein width. This indicates an actual build-up of tissue around the vein. These five species are Osmunda cinnamomea, Trichomanes boschianum, Woodsia obtusa, Dryopteris heragonop- tera, and Asplenium platyneuron. The leaf blade is thicker be- tween veins than it is at the veins in three species, namely Pteris vittata, Adiantum capillus-veneris, and Dryopteris marginalis. The measurements listed for LT in Table I are averages of those taken at the vascular bundles. There appears to be no great modification of leaf thickness due to environment as judged from leaves collected from the two sources. The averages of leaf thickness for these fern leaves was from 56 microns in Adiantum pedatum to 336 microns in Cyrtemium Jaleatum. INTERVASCULAR INTERVAL Measurements of IVI were taken where the adjacent vascular bundles were nearly parallel. The predominantly dichotomous branching of lateral vascular bundles sometimes occurs near the midvein as in Osmunda cinnamomea, Dryopteris hexagonoptera, and Nephrolepis eraltata. In Pteris vittata some lateral vascular bundles branch near the midvein and adjacent ones branch mid- way to the margin or very near the margin. Other ferns, such as Adiantum pedatum and Asplenium platyneuron, show several branchings of one vascular bundle. Ferns with the largest leaf segments have correspondingly large intervascular intervals. Table J shows that for Phlebodiwm aureum, Camptosorus rhizophyllus, Pteris cretica, and C yrtcemium falcatum the IVI is over 700 microns. Other ferns having inter- vascular intervals in this range, however, include plants with smaller leaf segments, such as Botrychiwm virgintianum, Tricho- manes boschianum, and Polypodium vulgare. 90 AMERICAN FERN JOURNAL Comparisons of IVI with the dissection of leaf (D) listed in the right hand column of Table J have been illustrated in Fig. J. Except for group 5, the IVI measurements for each group de- crease as the dissection of the leaf increases. In Fig. 1 the wide variations within each group are not related to the source. For example, the greatest IVI in group 1, Phlebodium aureum, and the smallest IVI in group 5, Adiantum capillus-veneris, were both grown in the greenhouse. It might be profitable to study the combination of segment size and leaf dissection as it may be related to intervascular interval. Vern Wiptn Vein width measurements show no constancy or correlation with source of material, genus, leaflet size, or leaf dissection. The range of vein width from 18 to 85 microns indicates that the tissue requirements of each species govern the size of veins. ANGLE OF VEIN BRANCHING In the genera Dryopteris and Polypodium the species studied show similar angles of branching of the veins. In Woodsia and Adiantum the two species of each genus which were examined have seca aagen different angles of branching. Woodsia obtusa shows and Woodsia scopulina 30° (not shown in the table). The esa are also very different, with angles of 32° and 48°. Since the angle of branching is measured from the midvein of the segment, the degree of dissection probably affects the angle at which lateral veins arise. The midveins in a highly dissected leaf are farther removed from the rachis. A correlation of the degree of dissection and the angle of branching, however, failed to show any significance for the species studied. A more detailed and refined treatment of these characteristics might be profitable. Discussion The tissues of the fern leaf do not show some of the specializa- tions commonly seen in the leaves of higher plants. The epidermis is not as highly specialized unless the inner lobes filled with chloro- plasts are considered to be a specialization. The mesophyll of VASCULARIZATION OF FERN LEAVES 91 many ferns is not noticeably differentiated into palisade and spongy layers as are many dicotyledonous leaves. This condition empha- sizes the horizontal continuity of the leaf tissues. The build-up of tissues around the veins of many ferns also allows for the greater distributive and collective area for the vascular bundle, as em- phasized by Armacost (1944). A 300 z a e o 2 a ie s | \ oe *| a FEC ECEREEEE : : 200 pi 3 Fabel H] Go| AY a os) A si Loe ‘a el Loa a ho ap ay 3 1 5 Po es md ° >a 10q } AG ad of) Ay oy ” on be cl Ho} Oo} +) > o| 2 = =e] oa] = ° Ae) Ea] Pal Ba) Bap OC} Hl <<] 2 Al 3 . BD. 0 Lobel Pale e222 252 Bs 44 4 4 55 ae Fig. 1. A HISTOGRAM OF CERTAIN COMMON FERNS ARRANGED IN GROUPS BASED ON THE DEGREE OF DISSECTION ILLUSTRATING THEIR RESPECTIVE INTER- VASCULAR INTERVALS MEASURED IN MICRONS. Although the average leaf thickness in this study of fern leaves is 149 microns, about one half of the species have a thickness between 60 and 140 microns. This is strikingly similar to the findings of Wylie (1946) in his study of ninety species of F lorida dicotyledons in which 53 species had an average thickness within this range, but the average thickness of all 90 species was 56 92 AMERICAN FERN JOURNAL microns—more than for these ferns. When compared with his study (Wylie, 1939) of 22 Iowa woody plants and 24 Iowa her- baceous plants, which had leaves averaging 152 and 163 microns respectively, these ferns are not significantly thinner. The factor which gives the impression of thinness in fern leaves is the absence of the abundant network of major veins that gives structural rigidity to many dicotyledonous leaves. The vein width has already been referred to in estimating the build-up of tissues around the vein. Although the veins in ferns are somewhat smaller at their tips and larger at the base of leaflets, many ferns fail to show, aside from a large midrib, the degree of difference referred to in other types of leaves as primary, second- ary, tertiary and so forth. This can be explained by one of the principles of Power (1923) which says that in most fern leaves there persists a distal dichotomy in the venation. The vein width does not correlate with the IVI, the angle of branching, or the degree of dissection of the leaves. It does correlate weakly with leaf thickness. Bower (1923) also suggested that in evolution the factors of leaf venation and leaf dissection do not advance together. If the method of Ettingshausen (1861) in measuring the angle of branching had been used, more consistent results might have been obtained and the angles would have been larger. Wylie (1939) mathematically defined the relationship between tissue organization and vein distribution in dicotyledonous leaves. He showed that with large ratios of spongy to palisade mesophyll the intervascular interval was large, and conversely, when there was a relatively thick palisade layer the intervascular interval was correspondingly small. Later Wylie (1948) pointed out the dominant role of the epidermis in two species of Adiantum. The lobed epidermal cells of ferns can be considered spongy mesophyll since they are parts of a horizontally contiguous tissue. In some ferns there is little or no mesophyll which may be considered to be vertically oriented palisade mesophyll. We should not be surprised, therefore, to find such large intervascular intervals in the ferns as compared with angiosperm leaves. In this study the average of all IVI measurements was 595 microns. For 194 species VASCULARIZATION OF FERN LEAVES 93 of dicotyledons from Iowa, Florida, the Pacifie coast, and New Zealand, Wylie (1939, 1946, 1954) found an average IVI of 143 microns. Philpott (1953) in a study of 47 species of Ficus found an average IVI of 155 microns. Lommasson (1948) in a study of 37 species of grasses found an average IVI of 141 microns. Since the IVI represents the distribution of the smallest veins of the leaf and since they are among the last tissues of the leaf to develop, they represent the final adjustment of the vascular tissue to the environmental conditions in which the plant is growing. These results again point to the often suggested idea that vein spacing is dependent upon the tissue organization of the leaf and that this in turn is governed by the demands of the constituent chlorophyll bearing cells. LITERATURE CITED ArMacost, R. R. 1944. The structure and functions of the border parenchyma vein-rib of certain dicotyledon leaves. Proc. Iowa Acad. Sci. 51: 157-169. Bower, F. O. 1923. The Ferns. Vol. 1. Cambridge ErTINGsHAUSEN, C. R. von. 1861. Die Blatt-Skelete der Dikotyledonen. ien LomMMASSON, R. C. 1948. Tissue relations of grass leaves. University of Iowa, Dissertation. eieveacies ANE, 1953. A blade tissue study of leaves of forty-seven species of us. Bot. Gaz. 115: 15-35. WYLIE, = ‘3. 1939. Relations between tissue organization and vein distribu- tion in dicotyledon leaves. Amer. J. Bot. 26: 219- 946. Relations between tissue organization and yaseulaticktion in br ices of certain tropical and Aha seer dicotyledons. Amer. J. Bot. 33: 721-726. —————. 1948. The dominant role of the epidermis in leaves of Adiantum. Amer. J. Bot. 35: 465-473. ———. 1954 Leaf prgriouomsgs of some woody dicotyledons from New Zealand. Amer. J. Bot. 41 191. University or Nepraska, LINCOLN, NEBRASKA 68508. 94 AMERICAN FERN JOURNAL Shorter Note A WroneL_y Locauizep SPECIES OF Pyrrosia.— Niphobolus cuneatus Kuhn (Bot. Zeit. 26:40. 1868) was described from material distributed by Hohenacker “in territorio rei publ. Ecuador” no. 5. Since Niphobolus was considered a synonym of Cyclophorus by Christensen, the new combination Cyclophorus cuneatus (Kuhn) C. Chr. (Ind. Fil. 198. 1905) was published, but apparently no pteridologist has ever looked carefully at the type since the original publication of the species. Both Nzphobolus and Cyclophorus are later names for the genus now known as Pyrrosia. In the “Index Vilicum,” there were only two American species recognized, both of them a query—C. americanus (Hook.) C. Chr. and C. cuneatus. The former was properly considered a different genus, Niphidium, by John Smith. And thus C. cuneatus remains as nominally the only American species of this genus. Through the courtesy of Dr. Eckhardt and Dr. Meyer, the type of '. cuneatus has been made available to me on loan. It is a minute specimen, containing about a half dozen fronds and a short length of rhizome. The material is sterile only. However, even in the absence of fertile fronds, it is readily identifiable as Pyrrosia rupestris (R. Brown) Ching, a species confined to Aus- tralia. The minute, obovate sterile blades, densely covered be- neath with stellate scales, these with a conspicuous, minute, round, red central disk and elongate straight white rays are characteristic; the rhizome scales agree also, these being elongate-lanceolate, light brown, and not ciliate. It is not at all likely that this Australian species occurs in Eeuador, and so a wrong label may be presumed. Hohenacker distributed plants from many parts of the world and a confusion of data is not at all improbable. The genus Pyrrosia is therefore to be excluded from the flora of the New World.—C. V. Morton, National Museum of Natural H istory, Washington, D.C. 20560. ReEcENT FERN LITERATURE 95 Recent Fern Literature THe GENUs ApDIANTUM IN CULTIVATION [POLYPODIACEAE], by Barbara Joe Hoshizaki. Baileya 17, nos. 3-4, pp. 97—191. f. 1—45. 1971. Published by the L. H. Bailey Hortorium, Cornell University, Ithaca, N. Y., $1.50.—In continuing her studies of cultivated ferns, Barbara Joe has given us a useful treatment of the maiden- hair ferns, which are popular with florists. These beautiful ferns are delicate, and do not like the hot dry atmosphere of most houses and apartments. Consequently, they are best grown in green- houses, where most of them are of relatively easy culture. Twenty species are recognized as being in cultivation in the United States, but almost all the named cultivars belong to Adiantum raddianum (usually called A. cuneatum), the “Delta Maidenhair.”’ The text is made most usable by the good line drawings of typical pinnules and by silhouettes of the blades. Many of the named forms are not easily determinable, because they vary greatly according to cultural conditions and also intergrade with each other extensively. It is indicated that some of the cultivars can arise repeatedly de novo from normal parents. This is evidently a genus that would repay genetical study..—C.V.M. POLLEN AND Spores or Cue, by Calvin J. Heusser. Pp. 1—95, pl. 1—60. 1971. Published by the University of Arizona Press, P.O. Box 3398, College Station, Tucson, Arizona 85700, $15.00. The present work describes, keys, and illustrates by photomicrographs 54 species of Pteridophyta of Chile, the Juan Fernandez Islands, and Easter Island [which is rather remote from Chile in its flora and more closely related floristically to Polynesia]. The work appears to be carefully done, using the Erdtman method of spore preparation. However, it may be doubted that this approach is truly valid at present. The scanning electron microscope is rev- olutionizing the study of spores, for it shows up many details ’ Mrs. Hoshizaki has asked me to point out two MH acmee 2 AR? Ge p. 104, “2-3 pi a gail is transposed with “3-4 pinnate’; on p. Aa” Is pinna, not a fror 96 AMERICAN FERN JOURNAL that can not be seen at all with the ordinary microscope, even with great magnification. The present photographs are hardly good enough for a clear identification of the species. The only out-and-out error that I notice is the inclusion of the description and illustration of the spores of “Davallia valdiviana,” although there is no such published species. Since the specimen concerned is deposited in the museum in Santiago it could have been properly identified. The genus Davallia does not grow in Chile, and so I would guess, without evidence, that the plant concerned is Rumohra adiantiformis; this species is related to Davallia, and although common in the Province of Valdivia is not otherwise included in this treatment. Since most of the text and illustrations in this work concern the phanerogams, the book will be of more use to students of the flowering plants.—C.V.M. F'Lora pE VENEZUELA, Vot. I. HELEcuHos, by Volkmar Vareschi. Instituto Botanico, Aptdo. 2156, Caracas, Venezuela. 1033 pp. 1968.—This volume, which is bound in two portions, illustrates well the difficulties of preparing detailed floras in tropical areas. Although about 700 species are treated, in some groups (e.g., Polypodium and the Cyatheaceae) the list of “critical species,” those that are published but which were not studied by the author, contains 60-80% as many as those accounted for in the flora! The volume is dated 1968 on the title page and 1969 on the soft- bound cover, but dates in the text indicate that the writing was completed as early as 1960. Critical recent literature is often not cited nor additions and corrections apparently made. Most sur- prising is the seriously flawed nomenclature. About 200 names are invalid for one or more reasons, including the lack of a Latin description, a basionym, or a type specimen. There are, however, 44 valid nomenclatural changes, 31 intentional and 13 uninten- tional. Of the former, 2 are illegitimate and 29 are legitimate; of the latter, 6 are illegitimate and 7 are legitimate. Reference is made to specialized details of anatomy, morphology, and paleo- botany, which is useful and missing from most floras. The illus- trations are well executed and helpful.—D.B.L. Exotic and Hardy Ferns Begonias BOLDUC’S GREENHILL NURSERY 2131 Vallejo Street St. Helena, Californie 94574 Open Saturdays and Sundays from 10 A.M. to 4 P.M. and by appointment Phone $63-2998—Area Code 797 Mail orders accepted TRIARCH INCORPORATED =A Service to Biologists = Te better serve the biological community: 1. TRIARCH produces and stocks nearly 3000 dif- ferent prepared microscope slide items for use in gen- eral botany, plant anatomy, plant physiology, phyto- pathology, bacteriology, general zoology, vertebrate histology, vertebrate embryology, and parasitology. 2. TRIARCH offers to prepare slides of new or spe- cial items for a nominal fee or in exchange for pre- served materials. 3. TRIARCH advertises in biological journals to help support the financial needs of the associated So- ciety. For our current catalog No. 15, or to request special service, write to: Paul L. Conant, President TRIARCH INCORPORATED Ripon, Wiseonsin 54971 Vot. 61 JULY-SEPTEMBER, 1971 No. 3 American Fern Journal A QUARTERLY DEVOTED TO FERNS Published by the MissoUR | RIOTANICAD AMERICAN FERN SOCIETY. a. ts =. S dooiael LIBRARY DAVID B. LELLINGER Cc. V. MORTON ROLLA M. TRYON IRA L. WIGGINS CONTENTS The Fern Vegetation of Aldabra Atoll................. F.R.Fossure 97 Leaf Epidermal Studies in Marsilea Joun T. MicKet aND Frank V. Votava 101 The American Species of Plagiogyria sect. Carinatae Davin B. Letuincer 110 The Genus Stenochlaena J. Smith with Description of a New Species R. E. Hourrum 119 in Dryopteris Taxa . WuitTier AND W. H. Waener, Jz. 123 The Gametophytes of Natural oa in the Fern Genus Pellaea Tuomas R. Pray 128 The Variation in Spore Size ging Germination Dea Schemochromic Blue Leaf-surfaces of Selaginella Denis L. Fox anp James R. Wetis 137 Shorter Notes: eS ne ES Gilbert, a Asiatic F 140 Recent Fern L’terature................. NMssaue! args . 142 DEC ¢ - 19/1 The American Fern Society Council for 1971 Warren H. WaGNER, JR., ene of Botany, University of es Ann Arbor, Michi: Joun T. Micke, New York Botanical Garden, Bronx Park, sioue oe York, 10458. Vice-President Ricnarp L. Havxe, Department of Botany, University of Rhode a Kingston, Rhode Island 02881 Secr LeRoy K. Hever, Division of Plants, Carnegie Museum, Pittsburgh, Penne sylvan 213. Tr go Davip B. Sepa Smithsonian Institution, Washington, D. C. cae Ciara preciety Representatives — en H, WacGner, JR., University of pi - © Q = Representative Roza M. Treen, me Barvant rivers S. Representative American oe Journal EDITORS Davin B. —— cee a Institution, Washington, D. C. 20560. C. V. Mo -. Smithsonian Institution, Washington, D. C. 20560. Rota } M. Tati, Jr. ray Herbarium, Harvard University, Cambridge, Mass. 0213 Tra L. od ns th ..Dudley Herbarium, Stanford University, Stanford, Calif , and published at 0 Elm ” Baltimore, Md. 21211 te postage paid Baltimore Orbe ase “of the Jour- nal are mem 0 wish to nges; membership lis terest, aining specimens or different localities. assist in obtain Matter rag — ter be addressed to the cage teettiaas ress, a for mem subscriptions, orders addressed to th hee tg bers = apm $5.50 gross, $5.00 net (agency fee $0.50); sent free to mem- $10. 00; 4 . mesa soct boy anys & — dues, $5.00; gprs —— ’ a xtracted repr ints red n vance will be I be furnished authors oy cost, plus to $6.25 each; single back numbers of 64 pages 0 ls, ag 65-80 pages, $2.00 each; ov ee pages, $2.50 each; Cumulative —_ olumes 1-25, 50 cents. Ten cant discount on orders of six volumes Dr. W. H. W: J ~ W. agner, Jr. t of Botany, University of Michigan, Ann Arbor, , Michlens 46104, is ri Librarian nd Guraor "Members may borrow books and the borrower pa: paying all shipping costs. spat Exchange Pew et ce er oye th Street, Seattle, —- 98115, : ion lists sent on Gifts and oo Z bac aay a “Herbarium specimens, bovanieal book | Siriaas. other = veleomed 8 American Fern Journal VoL. 61 JULY-SEPTEMBER, 1971 No. 3 The Fern Vegetation of Aldabra Atoll IF. R. Fossere! Aldabra is a ring-shaped coral island in the western Indian Ocean, 260 miles northwest of the northern tip of Madagascar and 400 miles east of Africa. Unlike the majority of coral atolls it has been elevated by tectonic forces about 5-6 m. The island is noted, for more than any other reason, as the last stronghold in the Old World of the giant tortoises that formerly roamed many regions of the earth. They now survive only in the Galapagos, where they are rare, and on Aldabra, where they are still abundant. The climate of Aldabra is dry and hot, and scarcely favors the development of much of a fern flora. Although ferns are prominent in several habitats, the fern flora is exceedingly poor. One species had been recorded prior to the Royal Society Expedition (1966— 1967) and, despite some weeks of active collecting by two botanists and several other scientists interested in the vegetation, the known fern flora still remained a single species, Acrostichum aureum L., until a tiny colony of Nephrolepis biserrata (Swartz) Schott was found by Mr. 8. A. Renvoise in a depression in a barren area of windbeaten rough limestone near the southeast coast. No further colonies, nor any other species, have been found on the atoll. That prominent and varied fern vegetation may be made up of essentially a single species is very odd, indeed. Acrostichum is a ' The observations reported here were ee while the author was a dog 4 of the Royal sents 8 Aldabra Expedition, in January and February, 1968. Thanks are due to the Royal Society sa & the Smithsonian Institution’ s Office of Memes nal Activities for the privilege of participating in this enterprise. Volume 61, No. 2 of the Journat, pp. 49-96, was issued June 24, 1971. AMERICAN FERN JOURNAL VoLuME 61, PLATE 17 “ +) + ¢ ORAL “CHAMPIGNON”’ wITH ACROSTICHUM AND TORTOISE. FIG. 2. ACROSTICHUM AUREU M PLANTS IN CORAL PITS NEAR THE BEACH. Photos Fic. Sma / by Jay Shaffer FERN VEGETATION OF ALDABRA ATOLL “99 big, coarse, leathery fern with erect.fronds. It is usually associated with mangrove swamps and coastal marsh vegetation throughout most of the Tropics. On Aldabra there are notable mangrove swamps, but Acrostichum was not seen in any of them. To vis- ualize where it does occur a brief account of the main surface features of the island is necessary. The island is a flat, almost complete ring of reef limestone, little higher than 7 m except for dunes, broken by several channels through which the tides race twice a day in and out of the great, shallow lagoon. The land area is about 60 square miles and, although entirely of limestone, exhibits several different types of surface. Much of it is extremely eroded into sharp pinnacles, ragged projections, deep pits, fissures, chimneys and crevasses. The edges are razor-sharp where the relief is too high for the tortoises to climb over it (Fig. 1, background), but where the relief is more moderate, the rubbing of their great shells has rounded off the projections (Fig. 1, foreground). This rough type of surface is locally called ‘““champignon’’; in Polynesia it is called “‘feo.”’ In the interior of some of the islands are areas of a flat, smooth or often slab-covered surface, called “platin,’’ with solution basins, fat soil areas, deep cracks and fissures, and numerous pools of water and mud. The pools have either sloping or vertical sides, and the water varies from fresh to as saline as sea-water or even more so. Storm beaches and occasional dunes of coral sand line some of the coasts, as on the northeast, east, and parts of the south sides of South Island, which is by far the largest of the four main sections of the ring of land. It is not surprising to find that luxuriant masses of Acrostichum entirely or partly fill many of the pools, especially and almost exclusively those with vertical sides and mud bottoms. It tolerates a remarkable range of salinities, by actual tests ranging from 1,700 to 15,700 parts per million dissolved salts. In these pools the masses of thick, creeping, branched rhizomes are dense, and the erect fronds are crowded in compact clumps which reach 1.5 or even 2 m in height from the mud. The larger pools could appro- priately be called marshes 100 AMERICAN FERN JOURNAL One sizeable area which I visited has a number of such pools, filled with Acrostichum, but in addition, several hectares of “platin” ‘Surface are intricately cracked. The cracks are filled with Acro- stichum. The rhizomes are mostly in the tops of the deep but narrow cracks; the abundant roots go down to the water as much as one meter below. The fronds here are smaller than in the pools, usually somewhat over 0.5 m tall, and the area has the appearance of an Acrostichum “meadow”. Scattered bushes of Ficus avi-avi L. make parts of this area look like a miniature fern “savanna.” Even more remarkable was an extensive area 100-300 meters wide of “champignon” extending along the east coast of the island back of the line of dunes (Fig. 1). In most of the island the “cham- Pignon” is covered by thick scrub, but here, possibly because of exposure most of the year to salt-spray blown by the tradewinds, there is little woody vegetation. Instead, in addition to a few herbs and dwarf shrubs hiding in fissures, pits, and chimneys, Acrost?- chum is the principal component of the vegetation (Fig. 2). Deep pits, a few centimeters in diameter, occur in great numbers in this limestone. Many of these have each an erect tuft of dwarfed Acrostichum fronds rising from a bit of rhizome down in the bottom of the pit. The fronds are from a few cm to 0.5 m tall, and even taller where they grow in chimneys or crevasses that afford better shelter. Occasional tiny plants grow in deep crevices. The whole area is very sparsely covered by these tufts of erect, green, leathery fronds. Many of the fronds have several terminal pinnae densely covered on the dorsal side by sporangia, seen under a lens to be arranged in a close network. Pits in rough limestone in almost any part of the island are likely to be occupied by Acrostichum, but along the east coast it is the principal vegetation on large areas. That one species of fern is able to assume such variations in habit and to tolerate such a range of conditions that it can fill a Series of niches normally occupied by other species shows the capabilities that may be brought out by a wide variety of habitats and little competition from other species. An interesting question 1s why Acrostichum is not found in Aldabra’s mangrove swamps, . which usually are regarded as its normal habitat. Perhaps there LEAF EPIDERMAL STUDIES IN MARSILEA 101 is merit in the suggestion that the mangrove fern is actually a species of Acrostichum other than A. aureum L. Watson (Malayan For. Recs. 6: 152-157. 1928) has distinguished A. speciosum Willd. as a species tolerating more frequent tidal inundation. No plants corresponding to the description of this, which has con- spicuously acuminate pinna tips, have been found in Aldabra. DEPARTMENT OF Botany, U. S. Nationa Museum, WasuH- INGTON, D. CG. 20560 Leaf Epidermal Studies in Marsilea JOHN T. MICKEL AND FRANK V. Votava! The aquatic fern genus Marsilea is widespread and well known throughout the world. Of the nearly 70 extant species, 20 are found in the New World. Many studies have been made on the genus from the standpoints of its physiology, morphology, anatomy, and development. Frequently there are difficulties in species identification since the taxonomic characters lie primarily in the reproductive structures—the sporocarps—and herbarium specimens are often sterile. A search was begun for vegetative characters to distinguish the species of North America, Mexico, and the West Indies. As part of this study leaf clearings were made, and some notable diversity was found within the group. Gupta (1957), in a preliminary study of the epidermal and soral characters of some American species of Marsilea, reported that the epidermis could be of some taxonomic value. Species differ, he said, in the walls of the epidermal cells (‘‘smooth, slightly wavy, and highly wavy”) and in the frequency of the stomata. He illustrated the epidermal patterns of 11 species and counted and * We wish to acknowledge the support of the National Science Foundation (Grant no. GB-13722 to the senior author). 102 AMERICAN FERN JOURNAL measured the stomata. His monograph of the Indian species (1962) reported differences in epidermal patterns under different ecological conditions. . () MARSILEA EPIDERMIS. Fic. 1. M. QUADRIFOLIA, UPPER. Fic. 2. SAME, LOWER. Fic. 3. M. MEXICANA, UPPER. Fic. 4. Same, Lower. Fic. 5. M. poy- CARPA, UPPER. Fic. 6. M. NASHII, UPPER. Our clearings of 13 species were studied in regard to stomatal size, frequency and ‘position, epidermal patterns, and certain ecological effects. Our findings differ somewhat from and go beyond those of Gupta sufficiently to merit reporting them here. Clearings were made of all 13 species found in North America, Mexico, and the West Indies. The specimens are cited in Table I, _ and all are in the herbarium of The New York Botanical Garden. -- Leaflets were cleared in 5% KOH for 1-2 days and were then LEAF EPIDERMAL STupDIEs IN MARSILEA 103 bleached in 20 % Chlorox solution. The clearings were stained with tannic acid and ferric chloride and mounted in diaphane. EpiIpERMAL PATTERNS We found roughly the same variation in epidermal patterns that Gupta reported: (i) those with long cells and smooth walls (Figs. 6,7), and (ii) those with wavy walls that range from only slightly wavy to strongly undulating (Figs. 8-15) ; but we could distinguish another group, those with short, blocky cells (Figs. 1-5). mooth walls are quite distinctive and are found in WM. nashii and M. tenuifolia. This character may be related to their very slender pinnae. Short, blocky epidermal cells were found caibistentiy in our specimens of M. quadrifclia, M. pclycarpa, and M. mexicana. Their walls vary in wall waviness, even within a species. In /. pelycarpa the cells tend to be more elongate than in the other two species. Most species have somewhat elongated cells with variously undulating walls (Figs. 8-15). There seems to be considerable variation, and we found it impossible to separate sharply those with slightly wavy walls from those with deeper undulations, as Gupta (1957) attempted to do. Marsilea mucronata, M. vestita, and. oligospora are generally considered closely related and have been combined by some authors (Hitchcock et al., 1969; Reed, 1954). Their combined range covers the western half of the United States. Gupta (1957) reported the latter two species to be distinct in epidermal pattern and stomatal frequency; however, we found all three to be essentially the same in these characters. In all species the cells toward the base of the pinnae have smoother walls than do the more distal ones; the former grade into the square or rectangular cells of the rachis epidermis. EPIDERMAL STREAKS We examined only aerial leaves and floating leaves, so we did not find the ecological differences in epidermal wall patterns of aerial versus submerged leaves reported by Gupta (1962) _ and Gaudet (1964a). The latter are said to have smoother cell 104 AMERICAN FERN JOURNAL walls than the aerial leaves. We found essentially no differences in epidermal cell walls between floating and aerial leaves. Van Cotthem’s drawings of M. quadrifolia (1970, fig. 585) show more. elongate cells with more deeply wavy walls than in our specimens. MaRsILEA UPPER EPIDERMIS. Fic. 7. M. rENUIFOLIA. Fic. 8. M. FOURNIER!. Fic. 9. M. wuncinata. Fic. 10. M. macropopa. Fic. 11. M. mouuis. Fie. 12. M. BERTEROI, The presence of stomata on both surfaces of his specimen suggests it was not floating and that perhaps in this case floating and aerial leaves display somewhat different epidermal patterns. One morphological feature which appears to be correlated with floating leaves is the presence of reddish streaks on the under _ surface of the leaflets. The presence of these streaks has been LEAF EPIDERMAL STUDIES IN MARSILEA 105 used occasionally as a key character in determining /. mexicana (Knobloch and Correll, 1962, p. 47). The streaks are also com- monly found in M. quadrifolia and M. pelycarpa. They are not merely chance streaks of pigmentation, but have an anatomical basis. In these areas the epidermal cells have thinner, smoother walls than do the surrounding epidermal cells (Fig. 4). The pres- ence of the streaks is strictly correlated with the absence of stomata on the lower surface, suggesting that these are floating leaves. Leaves with stomata on the lower surface have no streaks; an intermediate condition was seen in a single specimen of J. quadrifolia, Mcldenke 4608, which has a very low number of stomata and suggestions of streaks of thin-walled cells on the lower surface that. can be seen under the microscope. The streaks are by no means constant for any of the species mentioned above, but they are much more common in these than in any others. They have been seen in four specimens of the M. oligospora-vestita-mucronata complex from the western United States and in M. deflera from South America. This feature fre- quently occurs in species commonly found in water, which natur- ally have floating leaves frequently. The other species generally live in intermittent pools or moist meadows that are inundated only infrequently, and so only rarely have floating leaves. The fact that the cells have thin, smooth walls suggests that because of their submerged condition these areas are slow to differentiate and essentially remain in a juvenile condition. Whether these areas have a function is not known. STOMATAL FREQUENCY Stomatal size and relative frequency were determined for the upper and lower surfaces of all species. Frequency was meas- ured in five samples from each surface by counting the stomata visible in an ocular reticle (area 0.0715 mm2). The averages are given in Table I. As noted above, some plants have no stomata at all on the lower surface. It would appear from this feature, from the appearance of the plants (long petioles, few or no hairs), and rarely from label data, that these leaves were floating on the - 106 AMERICAN FERN JOURNAL water. Gaudet (1964b) found floating leaves of plants grown in culture to have no stomata on the lower epidermis. This is to be expected since the floating leaves of water lilies (Nymphaea) have stomata limited to the upper surface, and the submerged leaves of members of the Podostemaceae have no stomata on either surface. It is interesting to note, however, that totally sub- merged leaves of Marsilea will initiate stomata on the upper but not the lower surface (Gaudet, 1964a). MARSILEA UPPER EPIDERMIS. Fic. 13. M. MUCRONATA. Fig. 14. M. OLIGO- spora. Fig. 15. M. vestita. In spite of the location of the stomata on the upper surface and the frequent lack of stomata on the lower surface, the in- ternal anatomy of the leaves is of the traditional configuration, with a palisade layer under the upper epidermis and an open spongy mesophyll layer below. There are air spaces in the palisade layer directly beneath the stomata. Aerial leaves generally have an appreciable number of stomata on the lower surface, but the stomata are more abundant on the upper surface (see Table I). In only one specimen was the fre- quency greater below than above. Although the frequency of stomata on a single surface is fairly constant, it may vary from plant to plant in a Species, and the relative number of stomata on the upper and lower surfaces may also vary. Presumably this intraspecific variation is largely ecologically controlled. LEAF EpmerMAL Srupigs IN MARSILEA 107 Interspecific variation in stomatal frequency is quite pronounced in some cases. The number of stomata per unit area (0.0715 mm?) on the upper epidermis ranges from 13 to over 40, although most are below 30. Gupta (1957) noted that the species he examined could be distinguished in some cases on the basis of relative stomatal abundance. Marsilea vestita, for example, was reported to have three to four times as many stomata as many of the other species. Our data do not confirm this, for we did not find M. vestita to have an exceptionally large number of stomata. In fact, one specimen with larger than normal stomata had exceptionally few. Marsilea mexicana had more abundant stomata than any other species we examined. TaBLeE I. SromaTaL FREQUENCY AND S1zE IN MaARSILEA Frequency Size (Microns) Species & Collector Upper Lower Upper er BER slp fh . Braun—Puerto Rico acs et al. 8 18.2 178 Pp snaes 26.8 Sintenis ce 19.0 15.6 2t6 24.3 M. FOURNIERI C. Chr.— Mexico Brandegee, 5 Sept 1893 30.8 19.0 20.8 25.0 Brandegee, 16 Oct 1899 32.4 25.8 20.2 23.5 Carter & Moran 5394 20.8 18.0 21:1 24.2 Pringle 2434 8 10.8 27.9 29.6 i 5422 17.6 11.2 25.2 27.9 - MACROPODA Engelm.—Texas Poke 11246 20.2 16.0 25.2 pears Palmer 33856 12.8 14.4 0 See 26.8 M. MEXICANA A. Braun—Honduras & oe : Glassman 2054 34. 8 22.8 23:5 Pringle 2007. 37. Ps 0 20.8 eas M. MOLLIS Robins. & Fern.— Mexico ous os mn 604 22.8 12.4 eT eee Palmer 13 99:9 “Ho... 28 29.9 M. MUCRONATA os oo & Texas - Kolstad & Harms 29.4 17.8 24.9 29.2 Tharp, 30 May age 22.2 17.2 22.7 26.1 108 AMERICAN FERN JOURNAL TaBLe I. SromaTAL FREQUENCY AND S1zE IN MarstLeA (Continued) Frequency Size (Microns) Species & Collector Upper Lower Upper Lower M. NASHII Underw.— Bahamas Nash & Taylor 1411 22.4 14.0 30.2 34.6 M. OLIGOSPORA Ae —Idaho & Wyoming Nelson & Macbride 116. 26.2 18.2 21,9 28.0 Payson & Payson reais 24.4 21.0 22.6 26.4 Porter & Porter 9399 19.4 15.4 23.8 24.9 M. cmap binge & Grev.—Puerto Rico Britton et al. 67 26. 2 17.8 19.7 20.9 oaepatinta ea L.—Ontario, Mass. & Cult. Miller 6 647 29.4 0 2211 — Moldenke 4608 17.8 6.8 22.4 23.6 Seymour, Aug 1900 19.2 17.2 21.0 21.9 M. TENUIFOLIA Englem.—Texas Tharp, 11 Aug 1941 19.6 1726 26.5 29.1 NCINATA A. Braun—Texas Tharp, 3 Nov 1928 19,2 20.4. 26.5 M. eis Hook. & Grey.—Idaho, ca. & Utah Christ 1 9.6 26.8 28.8 Heller pe e > 18.2 25.1 26.4 Tillett 185 19,8 16.0 33.5 38, 2 STOMATAL SIZE Stomatal sizes were determined by measuring 20 stomata on each surface. Averages are shown in Table I. In every plant the stomata of the lower surface were found to be larger than those of the upper epidermis. Generally the difference is only 1-3 u, but occasionally it is more pronounced, as in /. uncinata and one specimen of M. oligospora. Certain species are outstanding in their very small stomata (M. polycarpa) or very large stomata (M. nashii), but most species have stomata 23-30 u long Within species there is aie hes relative dahorndt, -aithough considerable variation is seen in M. fournieri, M. mollis, and M. vestita. A polyploid series has bien reported in M. minuta of I ehra and Loyal, 1959), a species for which Gupta (1962) reports stomatal size differences. Lear EpipeERMAL SrupIgEs IN MARSILEA 109 ConcLUSION Our study has shown that considerable variation exists in the epidermis of Marsilea species, at least some of which has taxonomic significance. However, in order to determine more definitely the meaning of this variation, plants need to be grown under controlled conditions to determine the effects of water and light on the numbers and location of stomata; more extensive sampling is needed to establish the limits of variation within species; and more cytological information would be helpful in assessing the role that polyploidy plays in controlling stomatal size. LITERATURE CITED Gaupet, J. J. 1964a. Morphology of Marsilea vestita I. Ontogeny and morph- ology of the submerged and land forms of the juvenile leaves. Amer. J. Bot. 51: 495-502. . 1964b. Morphology of Marsilea vestita II. Morphology of adult and submerged leaves. Amer. = Bot. 51: 591-597. Gupta, K. M. 1957. Some American species of Marsilea with special reference to their epidermal and soral Sas Madrofio 14: 113-127. 1962. Marsilea. Botanical Monograph No. 2. Council of Scientific aad Industrial Research, New Delhi Hircucock, C. L., A. Cronquist, M. Ownsey and J. W. THompson. 1969. Vascular Plants of the Pacific Northwest, Pt. 1. Univ. of Washington Press, Seattle. ears I. W. and D. 8. Corre t. 1962. Ferns and — Allies of Chihua- a, Mexico. Texas Research Foundation, Ren MEurRa, P. N. and D. 8. Loyau. 1959. Cytological susie in Marsilea with particular reference to Marsilea minuta L. Res. Bull. Panjab Univ. n.s., 10: 357-374. Reep, C. F, 1954. Index Marsileata et Salviniata. Bol. Soc. Brot. II, 28: 1-61. Van Corruem, W. 1970. Comparative morphological study of the stomata in the Filicopsida. Bull. Jard. Bot. Nat. Belg. 40: 81-239 THe New York BoranicaL GARDEN, Bronx, NY 10458. 110 AMERICAN FERN JOURNAL The American Species of Plagiogyria sect. Carinatae! Davin B. LELLINGER? Plagiogyria is principally a genus of China and adjacent regions, and is usually said to have about a dozen species in the New World. These species and their nomenclature have never been adequately studied in the past, although Copeland (1929) attempted to distinguish them and named several new species. Ching (1958) has written an extensive treatment of the species found on the Asiatic mainland. Among ferns, Plagiogyria is unusual in having few or no tri- chomes at maturity. Young portions of the plants are covered with mucilage which is secreted by many, deciduous or incon- spicuous, glandular hairs. In the Asiatic species studied by Hennip- 968), the hairs are usually less than 1 mm long. Scales are absent, although patches of matted hairs and mucilage sometimes resemble the “amorphous” scales characteristic of a few species of Blechnum. Plagiogyria is unique among the leptosporangiate ferns in having sporangia with complete, oblique annuli and about 48 spores in each sporangium. The characters of the gametophyte are predominantly primitive, according to Stokey and Atkinson (1956). It is clearly an isolated genus, and in recent years has been put in its own family or order. The sterile fronds of the American species of sect. Carinatae, which supply all of the key characters, vary considerably within each species. The differences between the species are subtle, and immature or aberrant specimens may be difficult to place accu- rately; there seem to be no useful characters beyond those men- tioned in the keys. Because of the size overlap between extreme specimens of the various speciés and the general similarities of their fronds, descriptions are of little use, and so have not been included in this paper. ' chavs cost of publication of this paper has been borne by the Smithsonian nstit *I wish to thank the curators of 7 a from which I borrowed or examined specimens for their c coopera AMERICAN SPECIES OF PLAGIOGYRIA SECT. CARINATAE 111 Most of the New World species of Plagiogyria are quite re- stricted in distribution: one is from Hispaniola, one from Jamaica, one from Mexico, one from Costa Rica to Bolivia, and one from southeastern Brazil. The only widespread American species, P. semicordata, is found in Cuba and Jamaica and from Mexico southward to Venezuela and Bolivia. All of the species are ter- restrial at middle and upper altitudes in moist, cool mountains and are absent from coastal and interior lowlands. PLAGIOGYRIA sect. Carinatae Ching ex Lellinger, sect. nov estrial ferns up to ca. 1 m tall, with erect, dimorphic aa ti i crowded and persistent sere bases). Stipes and rhachises stramin- eous, brownish near the stipe base. Stipes 5-30 em long, usually triangular, suleate, only slightly expanded at the base; vascular bundles 3 at the base, fusing into a single V-shaped wel then an U-shaped trace, this becoming nearly ribbon-shaped in the rhachis; aerophores obsolete on the dorsal sfiarams of the ep bases or absent. Fronds devoid of scales, with mucilage-producing, glandu- lar hairs when young, deeply Pinnatifd: or the fertile sometimes pinnate), usually lanceolate, linear-lanceolate, or narrowly lan- ceate; rhachises terete below, suleate above. Sterile fide her- baceous or subcoriaceous; pinnae up to 15 em long, (3)5-10(12) * Until now the name Plagiogyria sect. Carinatae and the names of the sub- sections proposed by Ching (Acta Phytotax. Sinica 7; 135. 1958) have been invalid. Although they were in proper form and were provided with Latin diagnoses, the a of Art. 37 of the Code, which became diacties about five months before Ching published, were not ae I therefore wish to validate these cet by Saige: type species for them: PLAGIOGYRIA 8 ubsect. Adnatae Ching ex Lellinger, subsect. nov. TYPE SPECIEs: Lomaria adnata Blume fe Plagiogyria adncta (Blume) coe PLagioGyRia subsect. Euphlebiae Ching ex Lellinger, SPECIES: ieiiiaie scpkleetn Kunze [= Plogibayria eu} ‘hlebia oe dines “Mett. i ri sotto subsect. Pycnophyllae Ching ex Lellinger, subse:t. Typ. Es: Lomaria pycnophylla Kunze [= Plagiogyria sone tm ‘tons’ eal Ching also proposed sect. Euplagiogyrae, which is superfluous; a g to Art. 22 of the Code, the typical section must be an au shines ‘utonym Poa without citation of authority the name of the genus, i.e gogyr doubly so, the veins solitary or paired at the base and simple or - t 5). Fertile fronds herbaceous, generally longer and narrower and with longer stipes and shorter laminae than the sterile fronds; pinnae up to 12(15) em long, 2-4 mm wide, 7 SS eno ae Wg Bi ge Sa wees, We SSS ia eg Wit rg rt enh age baal Li DIAGRAMS OF VENATION TYPES IN THE AMERICAN SPECIES OF PLAGIOGYRIA sect. CaRinavak. Fic. 1. Simpe. Fig. 2. Parrep at THE BASE. Fig. 3. FORKED ABOVE THE BASE. Fic. 4. PAIRED AT THE BASE WITH ONE BRANCH FORKED ABOVE THE BASE. Fic. 5. PAIRED AT THE BASE WITH BOTH BRANCHES FORKED ABOVE THE BASE. the margins entire or irregular, often thin, somewhat involute, the veins 1-forked and sometimes uniting with an adjacent vein at the tip or forming a continuous commissure; receptacles somewhat enlarged and thickened; sori elongate; sporangia large, the stalk long, of 4-6 rows of cells, the capsule laterally compressed, the annulus oblique, continuous; spores 48 per sporangium, tetra- hedral, tuberculate to nearly smooth: z = ca. 25. Type specrEs: Plagiogyria argutissima Christ. AMERICAN SPECIES OF PLAGIOGYRIA SECT. CARINATAE 113 KEY TO THE AMERICAN SPECIES OF PLAGIOGYRIA SECT. CARINATAE Pinnae of sterile fronds 5-7(8) mm wide; veins simple and either solitary or paired at the costae, or veins simple and 1-forked (Figs. 1—3). Margins of sterile pinnae simply serrate, the veins beneath usually at QP. least aie prominulous (MEXICO) cs 26 AOC Ag ee pectinala Margins of sterile pinnae eran to doubly serrate, the veins beneath not ae an (BRAS) cer PON ak ed nga ey 3. P. fialhoi Pinnae of sterile fronds (7)8-12 mm wide; veins solitary and 1-forked or paired at the costae and 0-2-forked (Figs. 2-4). Sterile fronds decidedly narrowed at the base, the basal pinnae no more half as long as the longest pinnae. Sterile fronds 6-14(22) em wide,‘ the pinna margins usually biserrate, at least toward the apex (Jamaica, Cupa, Mexico to VENEZUELA = Bouivia). . P. semicordata Sterile pees not much narrowed at the base, the ar innae about aes ds as long as the longest pinnae. Sterile aaa 10—24(80) Sterile eke (10)13-24(30) em wide’ (Costa Rica a8 Bo.iviA). P. costaricensis Sterile fronds 10—-15(18) em wide. Sterile ae crenate-serrate mostly toward ies apex, sub- e toward the base; laminae mostly 15-18 em wide m4 P; ree AOA UH ye ee le ee eee maxonit Sterile ses sharply and evenly serrate throughout; laminae ostly 10-15 em wide (HispANIOLA)...... 6. P. serrulata 1. PLacioayria sEmicorpata (Presl) Christ, Farnkr. 176. 1897. maridium semicordatam Presi, Epim. Bot. ee 1849 [1851]. Type: Colom- bia, sin coll. (presumably PR or PRC not seen). Plagiogyria biserrata Mett. Abhandl. Senckenb. _ Naturforsch. Gesell., Frankfurt 2 [Farng. Plag. 8]: 272, t. 15, f. 1-18, 1858. LacroryPe: “Reg. alpina,’ Colonia Tovar, Venezuela, Moritz 400 (B not seen * Morton photo 10211; isolectotypes HBG not seen Morton photo 5339, US), chosen by Copeland (1929, p. 409). The other syntypes are Mexico, ae ky (B not seen) and ‘Hautes Andes de Truxillo et de Mérida,” ft, Linden 556 (B not seen; isosyntype BR Weatherby trash US) .4The latter specimen I identify as P. costaricensis. : Sterile fronds 6-8(12) em wide in South America, 8-12(15) cm wide in the West Indies, and 8-14 em wide in Central a with some specimens from Oaxaca and Morelos, Mexico, up to 22 cm 5 Sterile fronds 12-20 em wide in South Peter (9)12-15(30) em wide in a Rica 114 AMERICAN FERN JOURNAL Plagiogyria obtusa Copel. Phil. J. Sci. 38: 413. ¢. 13. 1929. Typn: Near Pico Turquino, Cuba, Leéa 11126 (US; isotype NY). RANGE AND HABITAT: Mountains of Cuba and Jamaica (alt 1650— 2220 m), of Mexico (Morelos and Puebia to Chiapas), Guatemala, Cesta Rica, and Panama (alt 2500-3150 m), and of Venezuela, Colombia, Ecuador, Peru, and Bolivia (alt 2500-3500 m), in moist, often shady piaces in coniferous or hardwood forests and subpéramo areas. Plajiogyria semicordata has biserrate pinnae that are narrower than those of P. costaricensis, which are broader and denticulate. The sterile pinnae of P. pectinata, in contrast, are simply serrate. ‘The apex of the sterile fronds of P. semicordata is more tapering than is that of P. costaricensis. It is possible that most of the South American specimens I have identified as P. semicordata— particularly those not from Venezuela—are small, contracted, and somewhat aberrant specimens of P. costaricensis, but I have no proof of this. np ED: Cuba: OrIENTE: Pico Turquino, alt 2000 m, Sezfriz 1091 (US), northern spur, alt 1850 m, Ekman 5465 (NY, US), alt 2040 m, Ekman 5496, 14546 (both US). Jamaica: Blue Mountain Peak, alt 2100-2225 m, Maxon 1413 (US), Underwood 2538 (NY), summit, alt 2100-2200 m, Maxon 9875 (NY, US); Between Portland Gap and Blue Mountain Peak, 400 ft, Wilson & Webster 623 (US); unlocalized, Gilbert 225 (NY), Hart 188 (US). Mexico: Veracruz: Orizaba, Miiller in 1855 (NY). Pursia: Honey Station, Pringle 15703 (F, US). Moretos: Zempoala, Lyonnet 1419 (US). Oaxaca: Cerro San Felipe, alt 3000 m, Conzatti 677 (US) ; Sierra de Clavellinas, alt 9000 ft, Pringle 4999 (NY, US). Cutapas: Cerro Hueytepec, 5km NW of Ciudad de Las Casas (San Cristébal), alt 8800 ft, Little & Sharp 9947 (US). Guatemala: HUEHUETENANGO: Cerro Canan4, between Nucapuxlac and Canan4, Sierra de los Cuchumatanes, alt. 2500-2800 m, Steyermark 49090 (F, US). Costa Rica: Cartaco: S slope of Voledn Turrialba, near the Finca del Voledn Turrialba, alt ca. 2000-2400 m, Standley 35133 (US); Behind the Hotel La Georgina, near Villa Mills, alt 2900 m, de la Sota 5100 (US), alt 3000 m, A. Jiménez M. 3401 (F). San José: Cerro de las Vueltas, alt 3000 m, Standley 43883 (US). Panama: Voledn Chiriqui, above El Potrero Camp, alt 2890-3025 m, Mazon 5334 (US). Venezuela: Aracua: Near Colonia Tovar, Fendler 325 (F, NY). Colombia: AnTIoguta: Hi Santa Rosa, alt 2500-2700 m, Lehmann (US). CunpI- NAMARCA: Macizo de Bogoté, Quebrada de Chicé, alt 2800-3000 m, Cuatre- AMERICAN SPECIES OF PLAGIOGYRIA SECT. CARINATAE 115 casas 6511 (US); Near La Gruta, Bogota, Little & Little 7848 (US); Guadalupe, m E of Bogota, Little & Little 9184 (US); Haught 5659 (F, NY, US). Ecuador: Azuay: Pdramo de Tinajillas, 30-50 km S of Cuenca, alt 11000-11500 ft, Camp E-2225 (F, NY, US). Peru: AMazonas: Summit of Puma-ureu SE of Chachapoyas, alt 3100-3200 m, Wurdack 1159 (F, NY, US). Bolivia: La Paz: North Yungas, Unduavi, alt 3500 m, Buchtien 2146 (US). 2. PLAGIOGYRIA pectinata (Liebm.) Lellinger, comb. nov. Lomaria pectinata Liebm. Vid. Selsk. V, 1: 233 [seors. 81]. 1849. Type: Cerro de Sempoaltepec, Oaxaca, Mexico, alt 8000-9000 ft, Liebmann (C not seen). Lomaria arguta Fée, Mém. Soc. Hist. Nat. Strasb. 5 [8 Mém.]: 70. 1857. Type: ‘Vallée d’Orizaba,”’ Cerro del Agua, Veracruz, Mexico, alt 2700m, Schaffner 98 (P? not seen). Plagiogyria sotenine Fourn. Mex. Pl. 1: 133. 1872. Type: Orizaba, Veracruz, Mexico, F. Miiller 723 (Hb. Lenormand not seen). The brief ease whick contrasts the simply serrate pinnae with P. biserrata Mett. (a synonym of P. semicordata), is sufficient to place the name. Pisin arguta (Fée) Copel. Phil. J. Sci. 38: 407. 1 Ra AND HABITAT: Hillsides and mountains of northern and central Niece (Chihuahua to Jalisco, Morelos, and Puebla), at 1750-3200 m altitude, in shaded, moist places in coniferous and coniferous-hardwood forests. Generally the basal pinnae of the sterile fronds of P. pectinata are about half as long as the longest pinnae, but in specimens from Oaxaca, Mexico, the sterile fronds have the lower pinnae even more reduced. ADDITIONAL SPECIMENS EXAMINED®: Mexico: Curnuanua: Near La Rocha NE slope of Sierra Mohinora, alt 7500 ft, Correll & Gentry 23112 (US); Mo- jarachic, Knobloch 5947 (US). Duranco: San Ramén, Palmer 124 (NY, US). Jaxisco: E of Mamantlan ca 15 mi SSE of Autldn by way of Chante, alt 8300 ft, Wilbur & Wilbur 2001 (US); near Etzatlan, Rose & Painter 7598 ap. 24a. Hipaueo: Beside the Rio Teponapa, vicinity of Zacualtipdn, a Moore 3207 (US). Pursia: Near Honey Station, alt 5800 ft, pai pie (F, NY, US). Morexos: Lagunas de Zempoala, Bro. Sanchez S. 466 (US). Mexico: La Juntas, Sultepec Road, alt 3200 m, Lyonnet 3119 (US) ; Zempoala, alt 3200 m, Matuda 28140 (US); Cerro de Acatitlin, alt 2000 m, Matuda 30762 ® The specimens (from Oaxaca south to Chiapas and — Guatemala) differ from typical P. pectinata in having the ov Fie of their sterile laminae reduced gradually below to 0.5-1 em long mere auricles; typical specimens (from Oaxaca north to Chihuahua). thee he lowest pinnae 0.35-0.65 as long as the longest ones 116 AMERICAN FERN JOURNAL (US) ; Crucero, 11 Nov 1934, Hinton 6823 (NY, US); Los Hornos, alt 2700 m, Hinton 2808 (NY); Nanchititla, Hinton 3600 (NY). Oaxaca: Sierra de Clave- llinas, alt 9000 ft, C. L. Smith 2063 (NY, US); Valley of the Rio Yelagago, ca. 20 mi. NE of Villa Alta, alt 6800-7200 ft, Mickel 929 (US); N slope of Cerro Peldn, along road from Ixtlan toward Tuxtepec, alt 9000 ft, Mickel 1142 (US). Catapas: Zonehuitz, Mtinch 102 (US); between San Cristébal las Casas and Tenejapa Center, alt 8400 ft, Breedlove 12907 (US); Barrio of Tuk, paraje of Matsab, Municipio of Tenejapa, alt 7500 ft, Breedlove 12565 (US). Guate- mala: HUEHUETENANGO: Top of Cerro Chemalito, Sierra de los Cuchuma- tanes, 3.5 mi. W of Santa Eulalia, alt 3100-3150 m, Steyermark 49927 (F, US). 3. PLAGIOGYRIA FIALHOI (Fée & era in Fée) Copeland, Univ. Calif. Publ. Bot. 19: 297. 1 Lomaria fialhoi Fée & Glaziou in vig ae . Vase. Brés. 1: 239, ¢. 7. f. 2 1869. Type: Serra os Orgéos, Est. Rio de Janeiro, Brazil, Glaziou 3326 (P not seen) DISTRIBUTION AND HABITAT: Southeastern Brazil, in the high portions of mountains, in low forests, according to £ehnem (1967). ADDITIONAL SPEC MENS EXAMINED: Brazil: Santa CaTARINA: Source of the Rio de Oratério, alt 1200 m, Smith & Reitz 10232 (US); Campo dos Padres between Fazenda Santo Antonio and Fazenda Campo dos Padres, alt 1400- 1650 m, Smith & Reitz 10377 (US). 4. PLAGIOGYRIA COSTARICENSIS Mett. ex Kuhn, Linnaea 36: 149. 1869 Plagiogyria anisodonta Copel. Phil. J. Sci. 38: 409, ¢. 9. 1929. Type: Dead crater of Voledn Poas, Prov. Alajuela, Costa Rica, alt 2800 m, O. Jiménez 1018 (US). Pararypss: Voledn Poas, Prov. Alajuela, Costa Rica, upper slopes between the Hotel and the crater, alt 2500-2640 m, Standley 34895 (US, , summit, alt 2644 m, Tonduz 10714 (US). Plagiogyria denticulata Copel. Phil. J. Sci. 38: 412, t. 12. 1929. Type: “Im ebelwald des Bergkammes der Laguna verde bei Comparapa [Com- arapa],” Depto. Santa Cruz, Bolivia, alt 2600 m, Herzog 1954 (US). Paratype: North Yungas, Bolivia, alt 3300 m, Buchtien 1908 (US). Plagiogyria latifolia Copel. Phil. J. Sci. 38: 411, t. 11. 1929. Type: Cani, 7 mi. of Mito, Depto. Hudnuco, Peru, alt ca. 8500 ft, Machride 3432 (US; isotype F). Type: Volcan Barba, Prov. Heredia, Costa Rica, Wendland 1066 (C? not seen) A drawing of the holotype (B) illustrates the broad, _ Coarse, sterile lamina and the fertile lamina only slightly reduced -at the base. AMERICAN SPECIES OF PLAGIOGYRIA SECT. CARINATAE 117 DISTRIBUTION AND HABITAT: Mountains of Costa Rica and the Andes, above (1200)1800 m altitude, in wet woods and shrubby paramos This species is larger and coarser than P. semicordata. The costae beneath often have what look like rather amorphous scales along them, but these are merely dried up patches of mucilage which have trapped numerous deciduous hairs. Sometimes a few cells from deciduous glands can also be seen; their terminal cells are swollen and reddish-brown. ADDITIONAL SPECIMENS EXAMINED: Costa Rica: ALAJUELA: Volcan Poas, near the summit at the edge of the crater, Burger 4352 (NY), upper lagoon, alt 2564 m, Pittier 2970 (US), Poas Lake, alt 8590 ft, Stork 2338 (US , along the road to the volcano, alt 6000-7000 ft, Scamman 7607 (US). Herepia: Vole4n Barba, alt 2700 m, Valerio 146 (US). Cartaco: La Pastora, near Voledn Irazu, alt 9000 ft, Stork 2378 (US,; N of Volcan Irazu, alt 10000 ft, Stork 1284 (NY); Voledin Turrialba, alt 3100 m, Torres R. 12 (US). Sin Joss: erro de las Vueltas, alt 2700-3000 m, Standley & Valerio 43540 (US), alt 3000 m, Pittier 10498 (US, ; Laguna de la Chonta, NE of Santa Marfa de Dota, alt 2000-2100 m, supa 42129 (US). Venezuela: Méinipa: Upper Andes of Truxillo and Mérida, Culata, alt 1600 toises [= 10240 ft], Denies 566 [a syntype of P. biserrcta Mett.] (B not seen; BR Weatherby photo US). ( olembia: Cunpinamanca: SE of Bogotd, alt 2800 m, Andre 976 (NY); Usaquén, near Bogotd, alt 2700 m, Uribe U. 208 (F); Quebrada de las Delicias, Macizo de Bogoté, alt 2650-2770 m, Cuatrecasas 5466 (F, US); Summit of Monserrate, E of Bogota, Little & Little 8189 (US); bart slopes of Péramo de Cruz Verde, alt 3150 m, Cuatrecasas 402 (US). : Headwaters of the Rio Tellez, above Soledad, alt 2900-3150 m, ce ei (US). Peru: HuAnuco: Yanano, alt ca 6000 ft, Macbride 3830 (F, US). Bolivia: La Paz: North Yungas, Unduavi, alt 3200 m, Buchtien 410 (F, NY), alt 3300 m, Buchtien 2156 (US), 2677 (F). 5. PLAGIOGYRIA MAXONII Copel. Phil. J. £ci. 38: 411, ¢. 17. 1929. Type; Summit of Blue Mountain Peak, Jamaica, alt 2100-2200 m, Mazon 9910 (US; isotypes F, NY). ' DIsTRIBUTION AND HABITAT: Mountains of Jamaica, in wet forest above 1700 m altitude. ADDITIONAL SPECIMENS EXAMINED: Jamaica: Sir Johns Peak, Watt 6 in 1910 (NY, US); Mossman’s Peak, upper eastern ridge and summit, alt 1700-1925 m, Mazon 9721 (F, NY, US); Blue Mountain Peak, alt 6500-7325 ft, Under- wood 1495 (NY), alt 7000-7423 ft, Mazon 1439 (US), Underwood 2554 (NY), alt 7420 ft, Harris 7144 (F). 118 AMERICAN FERN JOURNAL 6. PLagioGyRia serrulata (Willd.) Lellinger, comb. nov. Acrostichum serrulatum Willd. Sp. Pl. ed. 4, 5: 113. 1810. Based on Plate 81, Polypodium fuscum tenuissimis denticulis serratum, of Plumier’s ‘‘Tracta- tus de Filicibus Americanis.’’ Plumier’s plate illustrates several char- acteristics of this species (or of Plagiogyria in general): the evenly serrate pinnae with 1-forked veins, the acrostichoid fertile frond, and the sterile fronds scarcely narrowed below. His description mentions the blackish rhizome and the rhizome hairs, which are also characteristic of Plagio- gyria, but not of Polypodium, to which Plumier ascribed his plant. He gives the locality as “la bande du Sud au Grand Cul-de-sac, proche un “oxober 8 les ee appellent le fond épineux’”’ [Haiti]. Bl use in Urban, Symb. Antill. 7: 157. 1911, as “urbanz.” Tyre : Near Con sight Santo Domingo, alt 2200 m, von Tiirckheim 3408 (B not seen photo US, fragment US). Plagiogyria urbanti (Brause) Copel. Phil. J. Sci. 38: 413. 1 DISTRIBUTION AND HABITAT: Mountains of Hispaniola, in mossy and marshy places in pine forests above 1850 m altitude. ADDITIONAL SPECIMENS EXAMINED: Dominican Republic: Sierra de Ocoa, San José de Ocoa, Loma Sucia, alt ca 2300 m, Ekman H12002 (US); Cordillera Central, Moncidén, high ridge between the Rio Cerrobi and the Rio San Juan, alt ca 1900 m, Ekman H12790 (US); Sabana Nueva, between Piedra del Aguacate and Rio del Oro, alt 6200 ft, Howard & Howard 9123 (NY, US). Haiti: Massif de la Hotte, Torbec, top of Morne Formon, Ekman H7601 (NY, US). LITERATURE CITED CHING, es C. 1958. The fern genus ongspentins on the mainland of Asia. Acta Phytotax. Sinica 7: 105-154, t iia. CopELanp, E. B. 1929. The fern genus lca ca. Phil. J. Sci. 38: 377-417, 1-16. angie E. 1968. The mucilage ssteting ap on the young fronds of me leptosporangiate ferns. Blumea 16: 97-103. SEHNEM, "3 1967. Plagiogiridceas in R. Reitz, pit Ilustrada Catarinense, Parte I. Itajaf, Brazil. Sroxey, Ama G. — R. Arkinson. 1956. The gametophytes of iogyri ca (Bl) Mett. and P. semicordata (Pr.) Christ. vhctschint arg 6: 239-249. S. NATIONAL HERBARIUM, SMITHSONIAN INSTITUTION, Wasuineton, D. C. 20560. THE GENUS STENOCHLAENA 119 The Genus Stenochlaena J. Smith with Description of a New Species R. E. Hotrrum The genus Stenochlaena in Christensen’s “Index Filicum” (1905) included species belonging to three distinct genera. The three are superficially alike in having simply pinnate fronds with entire pinnae and free veins, the fertile fronds having narrow pinnae almost completely covered beneath with sporangia. But if one examines the vascular anatomy, scales and spores, one finds the superficial similarity hides a great diversity. The three genera are Stenochlaena s.str., Lomariopsis Fée, and Teratophyllum Mett. I studied these plants in the forests of Malaya, and wrote a paper describing their differences (Holttum, 1932). The differ- ences are also summarized by Copeland (1947), and the species of the three genera in the Malay Peninsula are described in Holttum, 1954. The genera Stenochlaena and Teratophyllum are confined to the Old World, but Lomariopsis is pantropic. ‘Thus all tropical American species formerly included in Stenochlaena belong to Lomariopsis (see Holttum, 1939), but Vareschi (1968) still places them in Stenochlaena, and his generic description includes a statement about vascular anatomy which applies to Stenocl.laena s.str. and is not true of the species he describes. In 1932 I expressed an opinion that Stenochlaena is related to Acrestichum (both genera have an exceedingly.complex vascular anatomy) and thus to Pteris, and I still think this possible; but C opeland has included Stenochlaena in the Blechnaceae. Lomaricpsis and Teratcphyllum appear to me to be related to Tectaria and Ctenitis. Six species of Stenocklaena are here recognized, two in Africa, the others in Asia, Malesia, and the Pacific. One of the latter has not yet received a valid name; as pointed out by Morton (1970) my attempt to give it one ignored the terms of the Inter- national Code of Botanical Nomenclature. In my paper of 1969 I also reported that examination of a type specimen in Presl’s herbarium necessitated a re-interpretation of his name S. ju, landi- 120 AMERICAN FERN JOURNAL folia and therefore a re-naming of specimens previously so called by me. As a comprehensive account of all species of the genus as now understood has never been published, I take this opportunity of providing one, to include the information of my paper of 1969. Only important synonyms and brief species descriptions are in- cluded. Further details may be found in the other literature cited. KEY TO THE SPECIES OF STENOCHLAENA Fertile fronds bipinnate, the sterile simply IRE SG RE ee args 1. S. tenutfolia Fertile _ sterile fronds both simply pinna nae commonly 15 pairs or more, ia eels pinnae to at least 15 X 3 em; fertile pinnae with very narrow sterile margin Seales on tip of rhizome narrow, spreading; eae of pinnae not ECHR ae eee ay 2. S. mildbraedii Seales on tip oi rhizome almost circular, appressed; pinnae articulate or n Lower vn at least, appearing fully articulate when dry; na-bases cuneate; fertile pinnae 2-3 mm _ wide; res evenly tuberculate throughout. . .3. S. palustris Pinnae not distinctly articulate; sterile pinnae with broadly _— to cordate bases; fertile to at least 5 mm wide; s with raised + broken ridges. Sterile ita ee broadly rounded at base, sometimes partly articulate; fertile eg 5 mm wide; spores with irregular broken es 4. S. milnet Sterile pinnae cordate at nae not at all articulate; fertile pinnae 7-10 mm wide; spores with parallel con- tinuous or broken ridges.......... 5. S. cumingtt Pinnae to 7 pairs; sterile pinnae to 10 X 2.2 em; fertile pinnae 3 mm wide with thin reflexed sterile margins 1 mm wide 6. S. areolaris — ‘ STENOCHLAENA TENUIFOLIA (Desv.) Moore, Gard. Chro 1856: 193; Morton, Amer. Fern Journ. 55: 165. 1965; 9; Tardieu in Humbert, Fl. Madag., Fam. 5, 1: 110. 1958. Lomaria tenuifolia Desv. Berlin Naturf. Freund. Mag. 5: 326. 1811. Rhizome to 12 mm diameter; scales on rhizome-tip narrow, spreading; sterile pinnae very variable in size, to 30 X 3 cm, not at all articulate to rachis; fertile pinnae to 15 em long with 10-12 pairs or more of pinnules ca. 5 em X 2m DistripuTion: East tropical to South Africa, Madagascar, Mauritius. THE GENUS STENOCHLAENA 121 Aaaeromenn: MILDBRAEDII Brause, Bot. Jahrb. Engler 53: 38 ; Tardieu in Flore du Cameroun 3: 353, pl. ee ie 5, 6. 1964. : Rhizome-scales elongate and spreading; stipe 100-150 em; frond 100-150 cm long; sterile pinnae 16—25 pairs, to 40 X 4 cm, not articulate to rachis; fertile pinnae 4-5 mm wide. Disrrispution: West Africa, Cameroons to Angola, eastwards to Uganda; climbing on Raphia palms. 3. STENOCHLAENA PALUSTRIS (Burm. fil.) Bedd. Ferns Brit. Ind. Suppl. 26. 1876. Handb. Ferns Brit. er 421. 1883. Holttum, Rev. Fl. Malaya 2: 412, fig. 241. 1954. Polyopdium palustre Burm. fil. Fl. Ind. 234. 1768. Lomaria ? juglandifolia Presl, Rel. Haenk. 52. 1825. Stenochlaena fraxinifolia Presl, Epim. Bot. 163. 1851. S. laurifolia Presl, ibid. 164 (see Holttum, 1969, pp. 14, 51). Rhizome long-creeping or climbing, ca. 5-7 mm diameter; stipes 15-20 em long containing up to 40 vascular strands; fronds 40-80 em long with 8-15 pairs pinnae; sterile pinnae articulate to rachis, very variable in shape and size, commonly ca. 15 X 3 cm, exceptionally to 24 em long but not proportionately wider, the base narrowly to broadly cuneate; fertile pinnae commonly 20 em long and 2-3 mm wide, with a very narrow, not reflexed, sterile margin; spores evenly tuberculate throughout. Disrripution: India through Malesia to N. E. Australia; in the Pacific to Samoa and Tonga. 4. STENOCHLAENA MILNEI Underwood, Bull. Torrey Bot. Club 33: 38. 1906. ° S. juglandifolia sensu Holttum, Gard. Bull. Str. Settl. 9: 139. 1937, non Presl ; Copel. Fern Fl. Philip. 428. 1960 Resembling S. palustris but sterile pinnae always broadly rounded at base and articulation to rachis not complete; stipe of large fronds containing a larger number of vascular strands; fertile pinnae ca. 5 mm wide; spores bearing irregular broken ridges. Type: Solomon Islands, Milne 518 (Ix). Disrrisution: Philippines, New Guinea, Solomon Islands. 122 AMERICAN FERN JOURNAL When I was in New Guinea in 1963, I noticed that this was the common species of Stenochlaena in the neighborhood of Lae, not S. palustris. It may be that the fern mentioned by Morton (1965, p. 166) as cultivated in the University of California Botanic Garden from spores collected in New Guinea is this species. Morton de- scribes it as a large and coarse vine; S. palustris has a more slender rhizome and smaller fronds, and I would not so describe it. 5. STENOCHLAENA Cumingii Holttum, sp. nov. S. laurifolia sensu Holtt. Gard. Bull. Str. Settl. 5: 259. 1932, non Presl; ibid. 9: 141. 1937; Copel. Fern Fl. Philip. 428. 1960; Holttum, 1969, pp. 1, 52. Stipes validus, basi fasces vasculares 100 vel ultra continens; pinnae steriles basi late cordatae, haud articulatae; pinnae fertiles 7-10 mm latae; sporae carinis + interruptis parallelis longitudi- nalibus ornatae. Type: Luzon, Cuming 226 (IK; isotype BM) Distripution: Moluccas, Philippines, New Guinea. This appears to me to be a very distinct species, though Copeland has expressed doubts. Stenochlaena milnei is almost exactly inter- mediate between S. palustris and S. cwmingii and may represent a hybrid. Cytological evidence is needed. 6. STENOCHLAENA AREOLARIS (Harr.) Copel. Philip. Journ. Sci. 2C: 406. 1908; Fern FI. Philip. 427. 1960. Lomaria areolaris Harrington, Journ. Linn. Soc. Bot. 16: 28. 1877. Rhizome slender; stipe of sterile frond ca. 15 em, of fertile ca. 30 cm long; sterile frond to 25 em long with 7 pairs of pinnae; pinnae to 10 X 2.2 em; fertile pinnae to 10 em long, 3-6 mm wide, with thin margins 1 mm wide reflexed at right angles. Disrrisvtion: Philippines, New Guinea, climbing on Pandanus. LireraTureE CiTep Cope.anp, E. B. 1947. Genera Filicum. Waltham, Mass. ottTtum, R. E. 1932. On Stenochlaena, Lomariopsis and Teratophyllum in the Malay Peninsula. Gard. Bull. Str. Settl. 5: 245-323. 1939. New species of Lomariopsis. Kew Bull. 1939: 613-627. _ 1954. A A Revised Flora of Malaya, II Ferns of Malaya. Govern- ment Printer, Singapore. SPORE SIZE AND GERMINATION IN DRYOPTERIS 123 Ho.rrvum, R. E. 1969. A commentary on some type specimens of ferns in the herbarium of K. B. Presl. Nov. Bot. Inst. Bot. Univ. Carol. Prag. 1968: 3-57. Morton, = oh 1966. Observations on cultivated ferns, VIII. Stenochlaena. r. Fern. J. 55: 164-166. —. 106. Review of ‘“‘A commentary on some type specimens of ferns in the herbarium of K. B. Presl.’’ Amer. Fern J. 60: 119-128. VaRESCHI, V. 1968. Flora de Venezuela, Vol. I. Instituto Botdnico, Caracas. Royat Boranic GARDENS, KEw, RicuMonp, Surrey, ENG- LAND. The epee in Spore Size and Germination Dryopteris Taxa DEAN P. WHITTIER AND W. H. WAGNER, JR.! Fern spores have provided a series of diagnostic characters for understanding the biology and relationships of the species. A number of studies have demonstrated, for example, that 32 spores in a sporangium are characteristic of apogamous species (Manton, 1950). The size of spores within a genus is commonly related to the ploidal level of the species (Blasdell, 1963; Wagner, 1966; Whittier, 1970). Sterile hybrids tend to have a greater variation in spore size than normal sexual species (Wagner & Chen, 1965; Kanamori, 1969). It also has been demonstrated that some spores of sterile hybrids can germinate and produce gametophytes (Morzenti, 1967; deBenedictus, 1969). Recently, Weinberg and Voeller (1969) reviewed external factors that control germination of fern spores and, among other observations, they found’ that spores from the same plants taken at different times gave very different rates of germination. * Research supported in part by NSF Grant oes Mr. Johannes Jensen prepared the spore materials for the experimen 124 AMERICAN FERN JOURNAL The woodferns, Dryopteris, present one of the most difficult of American fern complexes because of their tendency to hybridize. A number of workers have recently studied these plants cytologi- cally and have shown that many of the sexual taxa share one or more genomes with others. Except for D. fragrans, D. filiz-mas, and D. marginalis, all of the other sexual taxa have pairing affini- ties of the chromosomes with at least one other as shown in the following scheme (Wagner, Wagner, & Hagenah, 1969): Rr EE eT et ae Na etek es DD TP CODE Gere a ve be OO DDII ED AOI Ses Has na ew a we oP Pera ee aw es IISS TP a es I SSLL AP AMIGUIOMRN so he Ve O LL Be UMA es a ae A Ue eee Fy LLGG EE WOMOINE oO es os eee a ee re a GG DP, CUIOTAONG 6 ee ae is oe eis aah a SSLLGG Some of these, e.g., D. celsa, are evidently of hybrid origin, but they behave exactly like normal species because of their balanced chromosome complements and normal spore and gametophyte production. Sterile hybrids in Dryopteris, on the other hand, have usually one or more “unmatched” chromosome sets, and the resulting failure of these sets to pair normally produces spore abortion and sterility. The following investigation was undertaken to test the corre- lation of hybridity, spore size variation, and germinability in several Drycpteris species and hybrids grown under uniform conditions. MATERIALS AND METHODS The Dryopteris spores were obtained from plants in an experi- mental garden of the Matthaei Botanical Gardens at the Uni- versity of Michigan. Since the plants were grown under the same - conditions, environmentally produced variations in the spores or germination would be reduced to a minimum. In addition, the SPORE SIZE AND GERMINATION IN DRYOPTERIS 125 cytology of the plants has been studied and the chromosome pairing at meiosis is known. The spores were collected July 22, 1969, by the following method. A pinna was removed and was washed with water to remove the foreign spores. The washed pinna was placed in an envelope to dry and to release the spores. A sample (40 spores) was obtained from each taxon and measure- ments were made with a calibrated ocular micrometer. The stand- ard deviation (Li, 1964) was taken as a measure of the variation in spore sizes. The total length of the spore, including the peri- spore, was measured because the endospore can not be observed in spores of some hybrids (Wagner & Chen, 1965). The spores were sown on 2 ml of Knudson’s solution of mineral salts at pH 6.3 (Whittier, 1965) in capped, 1 dram vials. The cultural condi- tions included 12 hours of illumination (200 ft-c.) per day and a temperature of 24+ 1°C. Two hundred spores were observed seven days after sowing to determine the percentage of spore germination. RESULTS AND DISCUSSION Table I shows that the spores of the Dryopteris species had less variation in size, as measured by the standard deviation, than the spores of the Dryopteris hybrids. The germination of the spores of the Dryopteris species ranged from 78-95 % with one exception. Spores of one plant of D. celsa had 37.3% germination. Hybrid spores had low percentages of germination with two exceptions, 34.7% for D. celsa X marginalis and 36.0% for one plant of D. clintcniana X cristata. These Dryopteris taxa demonstrate that a greater variation in spore size, which is mainly produced by spore abortion, is correlated with the sterile hybrids of the genus. This supports the conclusion reached by others (Wagner & Chen, 1965; Kanamori, 1969). It is of interest to consider the taxa that had about 35% germi- nation. This level of germination was between the high percentages for the fertile species and the low percentages for the hybrids. The reduced germination for the spores of one plant of D. celsa probably was due to an internal factor which influenced the spores 126 AMERICAN FERN JOURNAL or spore development. It is unlikely that the reduced germination can be explained by environmental conditions because these spores were formed, collected, and grown under the same condi- tions as spores from other plants of D. celsa. The high percentage TaBLE I. Spore Size, STANDARD DEVIATION, AND PERCENT GERMINATION OF VARIOUS DryYoPTERIS TaXa. Taxa o Mean spore Standard Germination Dryopteris length in w deviation percent campyloptera 55.3 + 5.8 79.4 celsa 55.8 + 4.2 78.1 celsa 58.8 + 4.9 90.7 celsa 56.4 + 4.7 85.6 Isa 50.2 + 3.0 37.3 clintoniana 69.8 + 3.1 95.0 clintoniana 69.1 + 4.3 92.3 cristata 59.2 + 2.9 80.2 goldiana 44.0 + 2.4 82.7 intermedia 41.5 + 3.2 88.2 marginalis 44.6 + 3.5 90.9 marginalis si 3 + 3.5 89.3 campyloptera X a 48.8 +14.8 0.5 celsa X ets nian: 57.9 +19.6 0.5 celsa X gi 51.9 +13.6 1.5 celsa X m 55.3 +12.1 34.3 clintoniana X aia 56.2 +15.0 3.9 lintoniana X cristata 60.2 +10.1 36.0 ee X intermedia 51.5 +13.2 14 : X marginalis 52.9 +16.5 8.3 oon X spinulosa 57.5 +22.5 0.5 filix-mas X marginalis 61.1 +14.8 0.0 goldiana X marginalis 45.7 +12.0 0.5 of spore germination in the hybrids D. celsa X marginalis and . D. clintoniana X cristata was unexpected because low percentages of germination had been reported for sterile hybrids (Wagner & Chen, 1965). The reason for the high level of spore germina- tion for these two plants is unknown, but possibly it is due to a SPORE SIZE AND GERMINATION IN DRYOPTERIS 127 greater percentage of pseudomeiotic sporogenesis (Morzenti, 1962) in the sporangia of the plants. It should be noted that another hybrid, D. cristata X marginalis, also had a relatively high level of germination. Although in Dryopteris the percentage of spore germination is generally large for the species and small for the hybrids, the exceptions make spore germination by itself a poor character for suggesting hybrid origin for individual plants. LITERATURE CITED BLASDELL, = F. 1963. A pooh Oe, of the fern genus Cystopteris. m. Torrey Bot. Club 21 (4): ide semicvts, V. M. M. 1969. ne mixis in oo with special ete: to sterile hybrids. Doctoral Dissertation, Siasviaiies of Michig Kanamori, K. 1969. Studies on the sterility and size variation of oa in some species of Japanese Dryopteris. J. Jap. Bot. 44: 207-217. Li, J. C. R. 1964. Statistical cui Vol. I. Edwards Bros., Ann Arbor, Michigan. Manron, I. 1950. Problems of cytology and evolution in the Pteridophyta. Cambridge mbes Press, — ae idge Morzent1, V. M. 1962. A first report on pseudomeiotic sporogenesis, a type of spore reproduction by which “sterile” ferns produce gametophytes. Amer. Fern J. 52: 69-78. 1967. Asplenium plenum: a fern which suggests an unusual method of species formation. Amer. J. Bot. 54: 1061-1068 Wacner, W. H., Jr. 1966. New data on North American oak ferns, Gymno- carpium. Rhodora 68: 121-138. —————, and K. L. Cuen. 1965. Abortion of spores and ne as a tool in the detection of Dryopteris hybrids. Amer. Fern. J. 55: 9-29. , F.S. Waener and D. J. Hacenan. 1969. The log he (eydyteiis celsa) ao hybrids in Michigan—A preliminary report. Mich. Bot. 8: 137- WEINBERG, E. S. ee B. R. Vore.uer. 1969. External factors inducing germi- nation of fern spores. Amer. Fern J. 59: 153-1 Wuirtier, D. P. 1965. Obligate apogamy in Cheilanthes tomentosa and C. mishandieds. =~ Gas. 126: 267-2 81. 970. T in sexual and apogam- ous species of ferns. uke chias 20: 30-35. DEPARTMENT OF GENERAL BroLoGy, VANDERBILT UNIVERSITY, NASHVILLE, TENNESSEE 37203 AND DEPARTMENT OF Borany, University or Micutcan, ANN Arpor, Micuican 48104. 128 AMERICAN FERN JOURNAL The Gametophytes of Natural Hybrids in the Fern Genus Pellaea Tuomas R. Pray In the course of extensive field work relative to an investigation of Pellaea (Pray, 1968a, b, 1970) a number of collections were made on the western slopes of the Sierra Nevada in California of an undescribed member of the genus. Analysis of the plant sug- gests that apparently it is a hybrid (which will be described in detail elsewhere) of P. bridgesii and P. mucronata. The various features of the plant are mostly intermediate between those of the morphologically dissimilar parental species. As is to be expected in such a wide cross, the sporophyte is essentially sterile. At times, however, occasional individuals can be found to yield a few viable spores among the far more numerous nonfunctional products of abortive sporogenesis. Among the many individuals examined, two plants from a locality near Yosemite (Pray 1958, 1961) had sufficient functional sporangia to give a few spores to be sown on agar. In these sporangia, spore size was variable, with a few ob- viously larger than the rest. It was apparently the larger spores that were capable of germination. Accurate spore number per sporangium was not determined. It was, however, less than the normal 64 Spores were sown on nutrient agar as in previous investigations (Pray, 1968a). Most of the larger gametophytes were transferred to sterile soil 2 months after sowing to complete their development. The account which follows is necessarily incomplete due to the small number of gametophytes available. In order to determine whether the gametophytes could reach maturity and reproduce, most were allowed to continue growing, rather than be sacrificed for detailed microscopic examination. The usual complete develop- mental sequences were therefore not fully realized. Spores from the two hybrid plants used were sown separately. No differences were noted’ between the progenies, however; they are, therefore, de- scribed together. GAMETOPHYTES oF NATURAL Hyspribs IN PELLAEA 129 RESULTS The outstanding feature of the gametophytes from the hybrid plants was their extreme variability. None of these variations can be attributed to crowding or other unfavorable growth conditions. The germinating spore produces a filament as in other Pellaeas (Pray, 1968a, b, 1970). The filament may then give rise to a regular spathulate plate phase (Figs. 1, 2) which lacks a true apical cell. A few thalli (Fig. 2) may show a terminal cell with the appearance of an apical initial. Such a cell does not function as a true initial, however. Growth is generally diffuse at this phase. As the plate expands a marginal meristem with multiple initials is differen- tiated. This may occur in a terminal position. If so, the meristem soon occupies a shallow sinus and a typical cordate thallus with more or less equal wings is the result (F7g. 12). Although this form of thallus, derived from the most regular of the young plates, was observed in both groups of gametophytes, it is found in 1% or less of the total number of thalli. Sometimes a somewhat lateral meristem will differentiate (Figs. 5, 7). Strictly lateral meristems were also observed (Fig. 10). It is assumed that these latter types will eventually give rise to cordate thalli through the formation of a second wing and unequal growth rates of the regions lateral to the meristem. Thus the meristem is shifted to a terminal position. Asymmetrical thalli such as Fig. 17 are usually the de- rivatives of those with at least slightly lateral meristems at an earlier stage. Except for those relatively few gametophytes which followed fairly standard pathways of development for Pellaea, the re- mainder were quite aberrant. These usually lacked, or tended to lose, recognizable centers of growth. Frequently the shift from filamentous to two-dimensional growth failed to occur or was considerably delayed (Figs. 3, 4). This inability to initiate orga- nized two-dimensional growth could be permanent with the result that the filamentous phase continued to spectacular lengths. Figure 15 shows the major portion of a 2-month old gametophyte 4mm long. This individual has remained entirely uniseriate except for the vertical wall near the base. Some of its more regular con- AMERICAN FERN JOURNAL Vo.uME 61, PLatE 18 0 Oey, Lay Y CO te Sie Maret Ko) areteeay SAIN A) oa 3 ce i SS peaeees pec esaes 1 mm ‘Figs. 1-12. PELLAEA BRIDGES X P. MucRoNATA. Fics. 1-3, 5, 6, 911 FROM Pray 1861 (USC). Fias. 4, 7, 8, 12 rrom Pray 1858 (USC). GAMETOPHYTES OF NATURAL HyBriIDS IN -PELLABA 3 temporaries were typical cordate forms up to 4-5 mm in width. Sometimes the filamentous phase did eventually evolve non- terminal 2-dimensional regions (Fig. 4) which could differentiate .a more normal appearing plate (F7g. 6). In this instance a meristem might finally appear although such was not actually observed. Figure 6 might be interpreted as a case in which the filamentous portion is a secondary outgrowth and not the original terminal portion of the thallus. The precise developmental histories of ex- tremely peculiar, ameristic forms (Figs. 16, 17) is also uncertain. It is possible that these have arisen through localized expansion of median or basal areas of extended filaments. On the other hand. fairly regular early plate phases may have failed to differentiate meristems and developed filamentous outgrowths subsequently. It is worth noting that Figs. 4, 15-17, show much greater cell size than their counterparts. This suggests that continued cell enlarge- ment has occurred in the absence of normal rates of cell division. It is also possible, of course, that asa result of meiotic irregularities these particular thalli have higher chromosome numbers. Between the extremes represented by the above examples, there were intermediates of all sorts. Figure 8, with its large cell size, suggests a relationship to Fig. 4. Or perhaps this figure represents a form which has had more regular development and will ulti- mately organize a lateral meristem. Likewise, Fzg. 9, which lacks a meristem as yet, may be on the pathway to more normal growth or is becoming ameristic. From the original group of gametophytes, a very few grew on to become mature, normal-appearing thalli. I hese few were generally those whose development was most nearly normal. Thus among the survivors, a cordate thallus was sometimes encountered (Fig. 13). These tended to retain a rather shallow sinus. At the other extreme of variation were highly irregular, lobed forms (Fig. 14). The remainder were intermediate. Both types of game- tangia were produced and, at about one year, a few sporophytes appeared. These were apparently the products of fertilization by functional gametes. Several sporophytes survived for some time. They were rather similar to one another and more closely re- Puate 19 ? VotumE 61 = : 3 : é a GAMETOPHYTES or NATURAL HyBrips IN PELLAEA igs sembled the hybrid plants than they did either parental species. This lack of variability suggests apogamy. No evidence of asexual buds was noted however. All of the young sporophytes were eventually lost while they were still immature. Discussion It might be expected that in view of the intermediate nature of the hybrid sporophyte that the gametophytic generation would also show a compromise of the differences between the probable parents. The gametophytes of the parental species are markedly distinct in developmental features (Pray, 1968a). While the mature thallus of both species is cordate, the younger stages are distinguishable. In P. mucronata there is the regular differentiation of a strongly lateral meristem which only gradually becomes shifted to a terminal and eventually median position on the maturing thallus. In P. bridgesii, on the other hand, the marginal meristem is terminal from the beginning or at most only slightly asymmetrically situated; no truly lateral meristems occur. Thus it is the original site of the meristem which most obviously dis- tinguishes the gametophytes of the two species. In the hybrid, however, the great majority of the thalli never differentiate a meristem of any kind. Of those few which do, some are similar to one parent, some more like the other. This situation may indeed be a compromise between the parental pathways of development. It is of interest to note that certain Pellaea species, such as P. terni- folia, ave also characterized by just such variability in the site of meristem differentiation (Pray, 1968a). On the other hand, the filamentous or ameristic thalli are unlike anything shown by either parent. There is:also the question as to what extent the data from this particular hybrid are typical of nearly sterile fern hybrids. On this point, some observations on an other example are pertinent. A specimen was collected by the author in Arizona which appeared to be P. longimucronata (Pray 3218). More careful examination suggests that the plant is actually the hybrid of P. longimucronata and P. wrightiana, The specimen showed a high degree of sporangial VoutumeE 61, PLate 20 AMERICAN FERN JOURNAL aS SRI eee ee nieeat oh Imm i ‘Figs. 18-29. Peiuaka LONGIMUCRONATA X P. WRIGHTIANA FROM PRAY 3218 (USC). GAMETOPHYTES OF NATURAL HyBrips IN PELLAEA 135 abortion but did have a few apparently good spores. When the spores were sown on agar, a few gametophytes did develop which exhibited the same variability and high frequency of aberrant types (Figs. 18-29) as described here for the California plants. A few thalli (Figs. 18, 19, 23) had developmental sequences much like that of the parental species (Pray, 1968a). Filamentous (Figs. 20, 25, 26) and highly variable elongated forms (Figs. 21, 22, 24, 27-29) were also present. The great majority of thalli were thus quite different from what might be expected. The most obvious effect of hybridity was the extension of the filamentous phase and the inability to initiate normal two dimensional growth and marginal meristems. It would appear then that the preceeding account does not constitute an isolated instance, but rather that similar aberrant growth patterns are to be expected from the gametophytes of hybrids of this type. It may be significant that previous examples of gametophytes which closely resemble those of these nearly sterile hybrids have been recorded for certain apogamous ferns. Thus the thalli of some apogamous samples of P. andromedaefolia (Pray, 1968b) closely match the more peculiar forms of the hybrids described above. Other apogamous Pellaeas (Pray, 1970) may also have similar forms. On the other hand, that apogamy is not necessarily associated with aberrant gametophytic development is also clear. Again, in the case of P. andromedaefolia one apogamous popula- tion had quite regular gametophytes. Such exceptions were also noted in some other asexual Pellaeas, P. atrapurpurea for example (Pray, 1968a). In view of the resemblances between the hybrid gametophytes and those of many apogams, it is noteworthy that apogamy is often associated with taxa or “races’’ which have arisen through hybridization (Tryon, 1968). The difference is, of course, that these latter have become stabilized and self-per- petuating through apogamy, whereas the hybrids described in this paper apparently have not done so (as yet?). Finally, one might wonder whether the variations and irregulari- ties of development in the gametophyte generation of these hybrid ferns have the same basis as, or are analogous to, what Wagner 136 AMERICAN FERN JOURNAL (1962) has discussed in the case of the sporophytic features of hybrid ferns. For the hybrids described by Wagner there was frequent asymmetry and irregularity of various leaf characteristics by which the parental species differed. Particularly pertinent here was the conclusion that “hybrid expression is not a uniform and regular compromise.” While this statement also applies to the situation described in this paper for gametophytes, the causes may not be the same. In the sporophytes full genomes from the parental species are present and interacting. In the spores which produce the hybrid gametophytes such is probably not the case. Without a thorough investigation of sporogenesis and chromo- somal assortment in the hybrids, the chromosomal complements present in the various gametophytes remains unknown. It is, in fact, likely that they are variable chromosomally unless the functional spores are all the result of a non-reductional process. Chromosomal variability might explain the extreme variability of the gametophytes. LITERATURE CITED Pray, T. R. 1968a. The gametophytes of Pellaea section Pellaea: Dark- stiped series. Phytomorphology 18: 113-143. . Interpopulational variation in the gametophytes of Pellaea andromedaefolia. Amer. J. Bot. 55: 951-960. —————.. 1970. The gametophytes of Seog section Pellaea: Light-stiped ries. Phytomorphology 20: 137-1 Tryon, re F. 1968. Comparison es sexual a apogamous races in the fern genus sagen Rhodora 70; 1-24. Waaner, W. H., Jr. 1962. Irr sea morphological development in hybrid ferns. Moiuensieay i 12: 87-100. = 1068 DEPARTMENT OF BroLogicaL Sciences, UNIVERSITY OF SOUTH- ERN CALiForNIA, Los ANGELES, CA 90007. SCHEMOCHROMIC LEAF-SURFACES OF SELAGINELLA 137 Schemochromic Blue Leaf-surfaces of Selaginella Denis L. Fox! anp JAMES R. WELLS Schemochromic (i.e., optical or structural, non-pigmentary) colors are common in many animals, e.g., (1) Tyndall-blue scatter- ing in the iris of blue eyes; in skin areas of some birds, fishes, reptiles, and primates; in many feathers, as in jays; and (2) inter- ference-produced iridescence in pearl, nacre, wing-covers of some beetles, wing-scales of Morpho and many other butterflies, in feathers of peafowl, nummingbirds, pigeons, and other birds (Fox, 1953). Although plants excel animals. in the numbers and chemical varieties of true biochromes (colored molecules or pigments) that they store, instances of schemochromic coloration exhibited by plants are very rare. Moreover, the few examples involve no Tyndall-blue scattering, but are limited to the display of change- able spectral interference-colors, notably by a few seaweed species such as Iridophycus, Chondria, Zonaria, and others as viewed immersed in sea-water. A few exceptions found among land plants include some Lycopodium species? and certain Selaginella species, e.g., S. willdenovii (Desv.) Paker, a native of Malaysia.* The latter species has been introduced elsewhere, and is main- tained in greenhouses, including the University of Michigan’s Matthaei Botanical Garden, where the specimens studied were obtained through the kindness of Professor W. H. Wagner, Jr. ? Distinguished Scholar, es Institute of Pigexen 1970-1971; iam manent address: Division of Marine Biology, Scripps Institution of Oceanog raphy, University of California, a Diego; La Jol 4 CA 92037. I thank Dr. Warren L. Wittry, Director, and his colleagues at the Institute, for the honor i ces, such as Anemia makrinii Maxon; however, in this species at least thle disappears after drying —C.V.M. 138 AMERICAN FERN JOURNAL The upper surfaces of this plant’s leaflets exhibit, by sunlight or diffuse light, striking, changeable blue colors, reminiscent of the wings of Morpho butterflies but less brilliant. Microscopic inspection by reflected light, under medium powers, reveals count- less microspheric raised areoles, like masses of very minute beads, highlighted on the upper, projecting surfaces and exhibiting successive blue, blue-green, and purple or violet hues, depending on the increasing angle of vision. When viewed at a fairly acute angle, the leaf’s optical effects vanish, the surfaces then exhibit merely the green color of chlorophyll. It has been somewhat surprising not to encounter more pub- lished references to these striking changeable chromatic features over the upper leaflet surfaces of Selaginella. Walton and Alston (1938) commented upon the ‘metallic sheen of their leaves . . . due to the presence of particles in the cutin of the epidermis which reflect. certain rays of light.” While those authors came close to the factual basis of the observed coloration, the reference to ‘‘particles’’ was perhaps misleading, connoting as it does the presence of substantial, per- sistent material, perhaps of a solid character. The microsurfaces which reflect a changeable spectrum of optical colors are in fact air which is distributed into a pattern of minute films; hence the air is not discharged, nor the color quenched, by squeezing or rubbing a leaflet between the fingers. The optical colors disappear when the leaves are allowed to wilt. They are discharged also on wetting the fresh leaves briefly with either water or ethanol, but on evaporation or blotting of the fluid, the colors are restored. However, if a wetting agent, ¢.g., the clearing fluid Aerosol (Eimer and Amend) is applied, the immediate quenching of the bright colorsis permanent, even after the treated specimen is rinsed in water and superficially dried. Examination by transmitted light under high optical powers (400 X) reveals stomates o1 er the abavial leaf-surfaces and many small, light-refracting, air-filled spaces on the adaxial surfaces. The individual dark, light-refracting elements become suddenly clear as water gradually fills the air-spaces. ScHEMOCHROMIC LEAF-SURF! CES OF SELAGINELLA 139 The exposed cuticular layer of air-laden micro-spaces between the regular, projecting elements of the bead-like clusters, thus reminiscent of very minute interstitial soap-films, is the seat of the iridescent color-production. In much the same manner, entrapped microfilms of air give rise to the striking changeable blue colors of Morpho wings, which likewise are instantly quenched by the addition of foreign fluids having the same refractive index as the solid wing-protein, and which are restored on leaching away the fluid or allowing it to evaporate (Fox, 1953). Whether this unusual chromogenic manner of storing air on the upper surfaces of the leaflets may be merely an incidental condi- tion, evoked by photosynthetic, respirational and transpirational processes, and may be of no selective advantage to the plant, or whether it may in fact serve a role of physiological or biophysical importance (e.g., such as thermal buffering, light-screening, or water-retaining functions) must remain a problem for plant physiologists. LITERATURE CITED Fox, D. L. 1953. Animal Biochromes and Structural Colours. Cambridge University Press. Watton, J., Jn., and A. H. G. Atston. 1938. Chapter XVII: Lycopodiinae. ‘Fr. ‘Verdsor. ed. “Manual of Pteridology.”’ Martinus Nijhoff, The Hague CRANBROOK INSTITUTE OF SCIENCE, BLoomFiELD Hitxis, MI 140 AMERICAN FERN JOURNAL Shorter Notes ASPLENIUM KAMTCHATKANUM GILBERT, A Dusious EASTERN Asiatic Fern—lIn the “Index Filicum’” of 1905, Asplenium kamtchatkanum Gilbert (Fern Bull. 9: 54. 1901) is listed as dubious, with the comment “Athyrium sp.?” Apparently no one has ever examined the type, which is right here in the U. 8. National Museum. It was collected on the U. S. North Pacific Exploring Expedition on the Ochotsk Sea, Kamtchatka, by J. Small, be- tween 1853 and 1856, and had been originally identified as Asplen- ium filiz-femina L. The same species was collected also by the same expedition on Yezo Island. s species was overlooked in Komarov’s “Flora of Kamt- chatka” (1927), and in Hultén’s “Flora of Kamtchatka and Ad- jacent Islands” (1930) it is mentioned only as an unknown species of doubtful position. The type shows that this plant is the same as Athyrium pterorachis Christ, a species reported in Ohwi’s “Flora of Japan’ from Hokkaido, Honshu, Sakhalin, southern Kurile Islands, and Kamtchatka. It is a rather distinctive species, especially in the pubescent costae beneath, and clearly has nothing to do with Athyrium filix-femina. In fact, this species and some others have in recent years been removed entirely from Athyrium and placed in Dryoathyrium, Lunathyrium, or Parathyrium. If one assumes that the rachis structure is of fundamental importance in this group of ferns, then indeed this species and its allies will have to be separated from Athyrium, which differs in having the rhachis groove above interrupted and open to receive the grooves of the lateral pinnae. The matter has been discussed most re- cently by Hideaki Ohba in his paper “Considerations on the Genus Lunathyrium of Japan” (Sci. Rep. Yokosuka City Mus. 11: 48-55. 1965). He recognizes the single genus Lunathyrium Koidz. (1932) with three sections—sect. Lunathyrium, sect. Athyriopsis (Athyriopsis Ching, Acta Phytotax. Sinica 9: 63. 1964; Type: L. japonicum), and sect. Dryoathyrium (Dryoathyrium Ching, Bull. Fan Mem. Inst. Biol. Bot. 11:79. 1941; Type: L. boryanum; a synonym is Parathyrium Holttum, 1958). This seems the most SHORTER NOTES ; 141 reasonable treatment. The essential synonymy is as follows: LUNATHYRIUM PTERORACHIS (Christ) Kurata in Namegata & Kurata, Enum. Pterid. 309. 1961. [reprint without change of paging from ““Collection and Cultivation of Our Ferns and Fern Allies’? by T. Namegata]. Athyrium ier Christ, Bull. Herb. Boiss. 4: er “ge Asplenium kamtchatkanum Gilbert, Fern Bull. 9: 54. Asplenium ak Hultén, Kungl. Sv. Vet. ae ie 8(2): 237. 1930. Unintentional and illegitimate renaming of A. kamichatkanum Gilbert. Dry abe gh esitienteul Ching, Bull. Fan Mem. Inst. Biol. Bot. 11: 81. 1 (as ‘‘pterachis’’ erroneously). aera pterorachis Holttum, Kew Bull. 1958: 449. 1959. C. V. Morton, National Museum of Natural History, Washing- ton, DC 20560. A New Locauiry ror Psttotum NUDUM IN Sonora, MExico— In the spring of 1970, while studying the palm oases on the central coast of western Sonora, Mexico, I discovered a small colony of Psilotum nudum Palisot in Nacapule Canyon in the mountains near San Carlos Ray and about 10 miles northwest of Guaymas. The mouth of this canyon is located about 4.5 miles in from the coast, and the narrow canyon drains to the east out of steep and rugged volcanic mountains. The Psilotuwm grew at an alga-covered seep on the canyon wall in the shade of palms, Washingtonia ro- busta Wendl., and a few trees of a fig, Ficus padifclia H.B.K. While this locality is definitely in the Sonoran Desert; the shade of the canyon and the constant water supply provided by springs and seeps allows a number of plants with high water requirements to survive. Only one colony of Psilotum with about a dozen stems two to three inches high was found. A small collection of this plant (Boutin & Brandt 2814) is now growing in the Huntington Botanical Gardens Besides the Nicaoaie Canyon locality, three other stations are known for P. nudum in southern Arizona, Sonora, and western 142 AMERICAN FERN JOURNAL Chihuahua. It was recently reported in Syeamore Canyon, Arizona by Phillips (Amer. Fern J. 60: 30-32. 1970). It is also known in the Sierra Saguaribo of eastern Sonora in the drainage of the Rio Mayo, as reported by Gentry (Rio Mayo Plants, 1942), and from the Barranca del Cobre of Chihuahua, reported by Knobloch and Correll (Ferns and Fern Allies of Chihuahua, 1962) and also re- cently collected by Kimnach and Brandt (980). The habitat of Knobloch’s collection was not reported; however, each of the other collections has been recorded as being from a rock crevice or cliff site—Freprick C. Boutin, Huntington Botanical Gar- dens, San Marino, CA 91108. Recent Fern Literature THE CLASSIFICATION OF THE CYATHEACEAE, by R. M. Tryon, Jr. Contr. Gray Herb. 200: 4-53. 1970.—The scaly Cyatheaceae have mostly been divided into three traditional genera—Cyathea, with a complete indusium, Hemitelia, with a lateral, semispherical indusium, and Alsophila, with the indusium reduced to a small scale or absent. It has seemed to many that these three genera, although admittedly rather artificial, do represent natural groups, at least so far as the American species are concerned, for it has seemed that the species were more closely related to others in their own group than to those of other groups. However, Copeland, followed by Domin, did not agree and lumped all the species in a single large genus, without any subgeneric classification or other grouping. Recently, Holttum, working on Old World. species, established a new classification in which the scale characters were regarded as more important than the indusial characters. Tryon has followed Holttum by working on the New World species, and has developed a new system. He divides Cyathea into six genera—- Sphaeropteris, Alsophila, Nephelea, Trichipteris, Cyathea, and Cnemidaria. Sphaeropteris, mostly Old World, contains those species with the cells of the petiolar scales all alike. The other genera have the marginal cells variously differentiated. Alsophila and Nephelea have the petiolar scales with a dark apical seta, and REcENT FERN LITERATURE 143 sometimes also such setae on the back or margins. Alsophila, also mostly Old World, has no spines or, if these are present, they are not “‘corticinate”. Nephelea, strictly New World, has ‘‘squaminate”’ spines, the smaller ones being thin-marginate and _ scalelike. Cnemidaria, Cyathea, and Trichipteris have the petiolar scales lacking setae. In Cnemidaria, strictly American, the veins are areolate or, if free, the basal veins on each side are connivent to the sinus. Cyathea and Trichipteris have the veins all free and reaching the margin above the sinus. Trichipteris,! strictly Ameri- can, includes most of the exindusiate species that have been called Alsophila. Cyathea, also strictly American as here delimited, includes the indusiate species. In addition to the genera mentioned above, the generally recognized genera Lophosoria and Metarya are treated Tryon believes his classification to be natural. It may be so, although it-does have the appearance of being a ‘‘one-character” classification, the scale character being the decisive one in placing a species into a genus. It will certainly be useful in identifying specimens, and the stressing of the petiolar scale characters may influence collectors to be more careful to collect full specimens rather than detached pinnae. Tryon has not really justified his recognization of these groups as independent genera, his only statement being that he believes each genus to be a major evolu- tionary line, but that could and should be equally true also of subgenera. However that may be, Tryon is to be congratulated on a major contribution to the taxonomy of the Cyatheaceae. He is likely to be widely followed.—C.V.M. a correction rather than a different name it will continue to be credited to ; Pres! in 1822. The matter is exactly like the use of a wrong connecting vowel in specific epithets, which also must be corrected by Art. 73. 144 AMERICAN FERN JOURNAL “~ A Manvau or PLant Names (Second revised edition), by C. Chicheley Plowden. pp. 1-260. 1970. Published by Philosophical Library, 15 East 40 St., New York, N. Y., 10016. $10.00.—This book is intended for the gardener and amateur botanist who is interested in the meaning of plant names. Those treated are pri- marily plants cultivated for food, drugs, or ornament, and the commoner wild plants. The etymology of the Latin generic and specific names is given, and also a glossary of botanical terms used in descriptive botany. Needless to say, it will not replace a scholarly and indispensible work like R. W. Brown’s “‘Composition of Scientific Words” or B. D. Jackson’s “A Glossary of Botanic Terms,” but it may find a place by those interested in less technical and less complete works. I would not myself completely trust the etymologies given (although I must confess that I have not checked many of them). For instance, Glechoma is said to be derived from “glaukos” (gray), but it is surely from ‘‘Glechon,” the Greek name of some kind of pennyroyal. Pteris is said to be from “‘pteron, a wing,” but it is actually merely the Greek word pteris, applied to a fern; pteris may have come originally from ‘‘pteron,” which means primarily ‘feather’ rather than “wing.” Fragaria is not derived from “‘fragrans’’ but is a Linnaean variant of “fraga,” the Roman name for the strawberry; it may have ultimately been derived from ‘“‘fragrans”’ by the Romans. But no matter, the derivations are suggestive and doubtless correct in most instances —C. V. M. Exotic and Hardy Ferns Begonias BOLDUC’S GREENHILL NURSERY 2131 Vallejo Street St. Helena, California 94574 Open Saturdays and Sundays from 10 A.M. te 4 P.M. and by appointment Phone $63-2998—Area Code 707 Mail orders accepted TRIARCH INCORPORATED -A Service to Biologists = Te better serve the biological community: 1. TRIARCH produces and stocks nearly 3000 dif- ferent prepared microscope slide items for use in gen- eral botany, plant anatomy, plant physiology, phyto- pathology, bacteriology, general zoology, vertebrate histology, vertebrate embryology, and parasitology. 2. TRIARCH offers to prepare slides of new or spe- cial items for a nominal fee or in exchange for pre- served materi 3. TRIARCH advertises in biological journals to help support the financial needs of the associated So- ciety. For our current catalog No. 15, or to request special service, write to: Paul L. Cenant, President TRIARCH INCORPORATED Ripen, Wiseonsin 54971 Vou. 61 OctroBER-DEcEMBER, 1971 No. 4 American Fern Journal A QUARTERLY DEVOTED TO FERNS Published by the AMERICAN FERN SOCIETY ts DAVID B. LELLINGER Cc. V. MORTON ROLLA M. TRYON IRA L. WIGGINS is CONTENTS The Genus Grammatosorus..........-.0..- cee eeeeee C.V. Morron 145 The G Sel lla in Baja California, Mexico...... Tra L. Wieains 149 AMO &: Notes on the Ferns of Dominica and St. Vincent, II C. V. Morton anp Davin B. Letuincer 161 Two New Tree Ferns from Costa Rica............ Luis Dizco Gémez 166 Comparative Habitat Requirements for Spore Germination and Pro- thallial Growth of Three Ferns in Southeastern Michi Royce H. Hm 171 The Gametophyte of Thelypteris erubescens....Lenetre R. ArKinson 183 Shorter Notes: A paeomee ag of Isoétes in Louisiana; Jamesonia Ome PRORING os ok oh G hee ce a cas See ep tw oes wees 186 memes Wits Le in aa oe sea pnd ene ene 188 Subee to Voluaie 8425 os oo esc es 188 WM a aa i eile 191 Missoum! BoTramexr we Ty. ges _ JAN 10 1972 Che American Fern Society €ouncil for 1971 WarREN H. WaGner, JR., Poh gases of Botany, University of ag tee se Ann Arbor, Michigan 4 estden Jonn T. Mioxer , New York Botanical Garden, Bronx Park, _ Bons, Nes York. 10458. e-President Ricuarp L. Have, Department of Botany, University of atods — Kingston, sland 0288 Secretary LeRoy K. Henry, Division of Plants, Carnegie Museum, Pittsburgh, ae sylvania 15213. Tre — Davin B. Letuneer, Smithsonian Institution, Washington, D. C. Edttor- ie Cnied Hational Society Representatives Nn H. WaGner, JR., en of. _ chigan A.A.A.S, Representative ot M. Trron, JR., Harvard Uni A.I.B.S. Representative American pie Journal EDITORS Davip B. LELLINGER..... Smithsonian Institution, Washington, D. C. 20560. OC. V: oe oss: Smithsonian Institution, Washington, D. C. 20560. y He rbarium, Harvard University, Cambridge, Mass. 0213 Tra L. Wioains.. . Dudley Herbarium, Stanford University, Stanford, Calif An illustrated quarterly devoted to the general study of ferns, owned b b the American Fern Society, and publi at 3110 rave. Baltimore, Md. 21211. Sy reget postage paid at t Baltimore. The pages of the Jour- are open to me ro = wish to arrange exchanges; membership lists gs Bia me from different localities. assist interested in obtainin; Matter for r publication Gaul be addressed to the Editor-in-Chief. Changes plications for membership, subscriptions, orders Sn ae bee gross, net Bhs oct fee $0.50); sent free to mem- of the American Fern Soci (annual dues, $5.00; peer ati membership, $10.00; life membership, $100.00). Extrac on reprints, if ordered in advance, will be furnished a at cost, plus postage. : ; $5.00 to $6.25 each; — back numbers = less, $1.25; 65-80 pages, $2.00 each; over 80 pages, $2.50 each; ‘Cieeatien — to Volumes 1-25, 50 cents. Ten percent discount on orders of six volumes —— De We: EE ee of Botany, Unive: n, Ann Arbor, Atcha 4 ean ai, and Curato tor. gt i os ioe books and specimens sae tee tienes os ying all shipping costs. —. Spore Exchange , Mr.} D. Hall, 1225 Northeast 95th Street, Seattle, Washington 98115, is Director. Spores exchanged and lists sent on request. collection American Fern Fournal Vou. 61 OcTOBER—DECEMBER, 1971 No. 4 The Genus Grammatosorus C. V. Morton The almost unknown species Grammatosorus blumeanus Regel was published in the ‘Index Seminum” of the botanical garden in St. Petersburg in 1866; it was based on a cultivated plant that Regel had received from a man named Booth under the unpub- lished name of “Aspidium blumei.’”’ This Booth was probably John Booth, of Flottbeck, who according to the “Index Herbar- iorum Index of Collectors” collected in Java. Regel indicated that the original habitat was probably the East Indies. The connection with Blume is obscure. J. Milde obtained a specimen from Regel and studied it care- fully, and published his results in the “Botanische Zeitung” (26: 614-617. 1868). He concluded that the plant might represent either a sport developing in cultivation or a hybrid. This plant has remained essentially unknown for the last hundred years. It disappeared rapidly from cultivation, and no wild speci- mens resembling it have been discovered. Through the courtesy of the Director, Botanical Institute of the Academy of Sciences of the U.S.S.R. in Leningrad, I have been permitted to have the type on loan; a photograph of it is given herewith. It is a rather poor specimen, lacking a rhizome, The habit and venation are those of Tectaria. The general irregularity in cutting makes it appear that this may indeed be a hybrid, and this is partly con- firmed by the fact that most of the sporangia are abortive. A photograph of one of the larger and better developed spores made y the scanning electron microscope is shown in the accompanying figure, but even this is not a fully developed and regular spore. Volume 61, Number 3, of the JouRNAL, pp. 97-144, was issued Nov. 15, 1971. AMERICAN Fearn JoURNAL VouumE 61, PLate 21 Ho.otyre sHEet or GRAMMATOSORUS BLUMEANUS Rage. (LE) THE GENUS GRAMMATOSORUS 147 The chief peculiarity of Grammatosorus that caused Regel to describe this as a new genus is the sorus, which is strongly elongate along the reticulate veinlets; it is obviously indusiate throughout. Elongate, indusiate sori such as might occur in Diplazium are Scannina ELECTRON MICROSCOPE PHOTOGRAPH OF GRAMMATOSORUS BLU- MEANUS, X 70. not characteristic of Tectaria. There is however one species in which such sori occur, namely the extremely peculiar Tectaria dolichosora Copeland (Phil. J. Sci. 38: 136. 1929); in this the Sori are strongly elongate and indusiate; they curve around at the tip, thus becoming athyrioid in shape, and come back down the outer side of the veinlet, sometimes clear to the bottom, thus 148 AMERICAN FERN JOURNAL making a truly diplazioid, double sorus, with the indusia back to back and the sporangia spreading in both directions. In Grammat- osorus the sori do sometimes curve a little at. the apex where there is a vein fork, but they do not come back on the other side to form a diplazioid sorus. Other than in the sori there is no obvious alliance between Grammatosorus and 7’. dolichosora. In the latter the blade is much more divided and the axes are dark as in T. melanocaulon (Blume) Copel.! In Grammatosorus, the stipe and rhachis are stramineous. It may indeed be that Grammatosorus is a hybrid, in which case one of the parents might be Tectaria (Arcypteris) irregularis (K. B. Presl) Copel.; I would not dare guess as to the other parent, which would probably be in another genus. On the other hand, the existence of another species, T. dolichosora, with somewhat similar soriation indicates that this plant might well be a true Tectaria, for there is no evident reason to think that T. dolichosora is also a hybrid. In order to provide this plant with a nomen- claturally correct name, the following new combination is proposed: Aspidium blumei Kunze ex Mett. (1856), an entirely different plant. However, according to the Code both blumei and blumeanum are possible and correct, since thay are considered different names. Nationa Museum or Natura History, WASHINGTON, C 20560. ‘ This name is wrongly spelled ‘‘melanocaulis” by Copeland, apparently to make it feminine to agree with T'ectaria. However, the original melanocaulon of Blume is a substantive and therefore invariable. SELAGINELLA IN BaJA CALIFORNIA 149 The Genus Selaginella in Baja California, Mexico Ira L. WIGGINS The genus Selaginella was omitted from the ‘‘Vegetation and Flora of the Sonoran Desert’”’ (Shreve and Wiggins, 1964) because all material available to me at the time had been collected at altitudes above the elevation we had accepted as marking the upper limit of the Sonoran Desert, or came from areas in the Upper Sonoran Zone in northwestern Baja California. But when one considers the entire peninsula of Baja California, it contains seven species of Spike-moss, five of which Tryon places in the subgenus Selaginella section Tetragonostachys (A. Braun) Hieron. (Ann. Mo. Bot. Gard. 42: 1-99. 1955). The other two belong in the subgenus Stachygynandrum series Circinatae, according to Alston (in Ver- doorn, ‘‘Manual of Pteridology,”’ pp. 503-4. 1938). These seven Spike-mosses may be distinguished with the aid of the following key, and are illustrated here by line drawings in Plate 23 et seq. The drawings are offered as supplements to the excellent half-tone reproductions in Tryon’s paper in order to emphasize certain etails. KEY TO THE SPECIES OF SELAGINELLA IN Baga CALIFORNIA 1. Stems (with attached leaves) terete or quadrangular, not dorsiventrally flattened; plants erect, ascending, or decumbent but not forming rosettes, not rolling into a ball when dry; leaves narrowly lanceolate to linear-lanceo- late, with a median longitudinal groove on abaxial side (Subg. Selaginella). 2. Stems erect or ascending, not decumbent or prostrate; rhizophores produced only at or near bases of main stems (Ser. Arenzcolae). 3. S. bigelovit 2. Stems creeping or prostrate, the main ones neither — nor . strongly ascending; rhizophores along lower side of most of s 3. Setae at apex of vegetative leaves and tk denticulate scabrous; leaves markedly glaucous; sporophylls acumina CRae ea EO OR oo cs epee Ox} 4 ae Hr ne em ees 2. S. asprella 3. Setae at apex of leaves smooth, filiform and ephemeral, or lacking; leaves bright green or persistently ashy-gray (in S. cinerascens), not glaucous. VotuME 61, PLATE 22 AMERICAN FERN JOURNAL No’ / a A a4 Q "4 ‘Qe . —s “a » A A A 0 ( ’ ’ : 25° ‘ DisTRIBUTION OF SELAGINELLA IN Baga CaLiroRNIA. SOLID CIRCLE = ARIZONICA; OPEN CIRCLE = 8. ASPRELLA; OPEN TRIANGLE = S. BIGELOVII; OLID TRIANGLE = 8. CINERASCENS; OPEN SQUARE = S. EREMOPHILA; SOLI SQUARE = S. LEPIDOPHYLLA; Souip sTaR = S. PALLESCENS. SELAGINELLA IN BAJA CALIFORNIA 151 4. Ultimate branchlets mostly 2-4 mm long, these and main stems iene aaa leaves ashy-gray, their tips acute or sometim cous, not setiferous; plants growing on soil, not in eth crevices (Ser. Sartoriz)....... . S. cinerascens 4. Ultimate branches mostly 5-12 mm long; tips of branches green, not ashy, their tips bearing straight, slender, more or less deciduous setae, or the terminal seta (in S. eremophila) filiform and ephemeral, usually visible only on leaves at tips of branchlets (Ser. Eremophilae). 5. Microspores usually orange; terminal seta straight, milk-white, 0.1-0.3 mm long, never Ss leaf blade, Often. deciduous in are. <2. <6 oan ees . S. arizonica 5. Microspores usually yellow; terminal me filamentous, tortuous, often one-half to equaling length of blade, deciduous very pa and obvious only on leaves at tips Of prawiir branchlets..). 22-.+.6s e2 ne er 3 5. S. eremophila 1. Stems and branches flattened dorsiventrally; plants prostrate and rosu- late when moist, branches curling inward and upward to form a compact ball when dry; leaves dimorphic, those on upper side of stem smaller than ones on lower side, neither series longitudinally grooved (Subg. Stachy- gynandrum Ser. Circinatae). Leaves on lower side of stem suborbicular, rounded at apex, reddish or brownish; leaves on upper side eS Ave broadly bay without a terminal seta, conspicuously white-margined......... 6. S. lepidophylla Leaves on lower side of stem ovate, pei suborbicular, silvery white, acute and short-setiferous at apex; leaves on upper side of stem ovate to orbicular, bearing a stout, terminal, denticulate-scabrous seta, less MAArG@dIe wWhithnaremens, 5% 25-62 y- se oes eda es ne 7. S. pallescens Descriptions published by Maxon (Smiths. Mise. Coll. 72(5): 1-10, pls. 1-6. 1920) and by Weatherby (Amer. Fern J. 33: 113- 119. 1943; J. Arn. Arb. 25: 407-419, pls. 1, 2. 1944) admirably cover the species within the subgenus Selaginella that are known to occur in Raja California. Likewise, those in Correll’s “Ferns and Fern Allies of Texas,” pp. 28-43. 1956, and in Knobloch and Correll’s “Ferns and Fern Allies of Chihuahua,” pp. 16-32. 1962, deal with some of the species in subgenus Selaginella and both of the species in subgenus Stachygynandrum. Therefore, I include ere only citations of specimens from Paja California examined, together with comments on habitats and distribution. Vo.tuME 61, PLATE 23 AMERICAN FERN JOURNAL SELAGINELLA IN Baga CALIFORNIA 153 The line drawings were made with the aid of a Spencer camera lucida and various lens combinations of a Rausch and Lomb compound microscope. Abbreviations of herbaria from which specimens were studied are those recommended by Lanjouw and Stafleu in “Index Her- bariorum,” ed. 5, 1964. 1. SELAGINELLA ARIZONICA Maxon, Smiths. Misc. Coll. 72(5): 5, pl. 3. 1920. On rock faces, in crevices in and between boulders and ledges, often on soil or gravel at the bases of boulders, but rarely away from boulders or cliff faces, in Gila, Graham, Maricopa, Pima, and Pinal counties, Arizona, east in southwestern Texas (although not known to occur in New Mexico), south into northern Sonora, and known from a single collection near Loreto, Baja California. Mostly at aie is as m alt. EXAMINED: Baja California: Arroyo Undo [Hondo], [near] St a ae 930, M. E. Jones (DS). The SS distribution of this Selaginella is puzzling, for there are areas along the eastern slopes of the Sierra San Pedro Martir and other ranges in Baja California that would appear to provide suitable habitats for it. More intensive search may reveal it in such areas. 2. SELAGINELLA ASPRELLA Maxon, Smiths. Misc. Coll. 72(5): 6, On rock faces, in crevices on ledges, at bases of boulders, cliffs, and around talus, Los Angeles, Orange, and San Bernardino Figs. a-e, SELAGINELLA ARIZONICA: 4 = FUNCTIONAL LEAF FROM UNDER SIDE OF DECUMBENT STEM, THE TERMINAL SETA INTACT; b = LEAF FROM SAME AREA, SETA DETACHED; C = LEAF FROM UPPER SIDE OF STEM AT SAME LEVEL AS & AND b, TERMINAL SETA DETACHED; d = MATURE MICROSPORE; e = MEGA- SPOROPHYLL, ABAXIAL SIDE. Fics. f-i, SELAGINELLA ASPRELLA: f = LEAF FROM fo) STEM; g OPPOSITE f; h = MEGASPOROPHYLL, ABAXIAL SURFACE; ATURE MICRO- SPORE. Figs. k-p, SELAGINELLA BIGELOVII: k = MATURE peony OUTER SURFACE; 1 = MIcROSPORE, COMMISSURAL SIDE; m = MICROSPORE, OUTER SURFACE; n, 0 = LEAVES FROM SAME LEVEL ON OPPOSITE SIDES OF ASCENDING STEM; p = ABAXIAL VIEW OF MEGASPOROPHYLL. 154 AMERICAN FERN JOURNAL counties, California, south along the eastern escarpment of the San Jacinto and Cuyamaca Mountains into Baja California. SPECIMENS EXAMINED: Baja California: In crevices of granitic rocks along streams, near La Encantada, Sierra San Pedro Martir, alt 7800 ft, 22 Sept 1930, Wiggins & Demaree 5002 (DS); Rocky cliff in first canyon off Cafion del Diablo on NW side of Picacho del Diablo, eastern flank of Sierra San Pedro Martir, alt 6000 ft, 15 June 1954, Chambers 643 (DS); Rim overlooking cafion between main range and Picacho del Diablo (Picacho de la Providencia), near Lat. 31° N, Long. 115° 24’ W, about 9000 ft, 5 Sept 1961, Wiggins 16635 (DS); Crevices in granitic rocks on SE slopes, Cerro Venado Blanco, Sierra San Pedro Martir, 15 Sept 1968, Moran 15670 (CAS). This species is far more common among igneous rocks and cliffs in the Sierra San Pedro Martir than the small number of specimens in herbaria would indicate. 3. SELAGINELLA BIGELOVII Underw. Bull. Torrey Bot. Club 25: On dry, sunny, rocky slopes and ridges or on soil in partial shade of sparse stands of chaparral, often abundant around boulders or on steeply sloping to nearly vertical cliffs and rock faces, inner Coast Ranges, Santa Clara County, California, southward to the mountains of the Cape Region, Baja California, at elevations between sea level and 1500 feet in the northern part of its range, and upward to over 6000 feet in the Cape Region. SPECIMENS EXAMINED: Baja California del Norte: Brush-covered hills SW of Tijuana on Agua Caliente road, 24 Feb 1934, R. S. Ferris 8470 (DS); South facing slope about 11.4 mi E of Rodriguez Dam, east of Tijuana, 24 Jan 1966, Wiggins 20373 (DS); Near Rodriguez Dam, 10 mi E of Tijuana, in tufts at bases of rocks, 1 Apr 1947, H. F. Copeland (DS); Hillsides NE of Ensenada, alt 200-400 ft, 23 Feb 1930, Wiggins 4212 (CAS, DS); About rocks on hill- sides facing the sea, Ensenada, 7 Apr 1921, I. M. Johnston 3005 (CAS); San Carlos River (S of Ensenada), 10 Sept 1923, Eastwood 12391 (CAS); San Car- los canyon, 10.6 mi E of Ensenada-Santo Toms road, 8 Apr 1954, Wiggins Chapala road, along track to Cerro Blanco and San José, Lat. 29° 42’ N, Long. 114° 25’ W, alt ca 1500 ft, 7 Feb 1962, Wiggins & Thomas 120 (DS); SELAGINELLA IN BaJA CALIFORNIA 155 Catavifid, 14 Apr 1954, Dorothy R. Harvey (CAS); North-facing rocky wall of canyon, 2 mi N of Mision de San Borja, 16 May 1959, I. L. & D. B. Wiggins 14843 (CAS, DS); Rocky ridge above Rancho Aguajito, 32.5 mi SE of El Rosario, 15 May 1956, Howell 31012 (CAS); Same area, Porter 162 (CAS); Rocky talus slopes 5 mi W of Mision de San Borja, along road to Rosarito, alt ca 1000 ft, 18 Feb 1962, J. L. & D. B. Wiggins 16739 (DS); Between boulders in arroyo bottom opposite store at El Arenoso, alt ca 2500 ft, 21 Jan 1960, Porter 162 (DS); On exposed rocks E slope of Cerro Barranco, Sierra Santa Lucia, near Lat. 26° 56’ N, Long. 112° 28’ W, alt ca 1300 m, 10 Mar 1964, Moran 11784 (DS); Locally common on north slopes near summit, Cerro San Juan, near Lat. 27° 58’ N, Long. 113° W, alt ca 1200 m, 4 Feb io Moran 11597 (DS); NW slope, Cerro de la Mina de San Juan, near Lat. 2 cOING Long. 113° 38’ W, 25 Mar 1960, Moran 8051 (DS). Territorio Baja California del Sur: Rocky mountain top, in oak forest, Cerro de la Giganta, alt 5000 ft plus, 1 Mar 1939, Gentry 4274 (DS); On steep, north-facing ane on ridge NW of main peak of Cerro de la Giganta, alt 1200-1350 m, Lat. 26° 8’ N, Long. 111° 36’ W, 25 Nov 1947, Carter, Alerander & Kelli 2050 (DS); Among granite rocks at summit, in Quercus idonea with occasional Q. devia and Pinus cembroides, Sierra de la Laguna E of Todos Santos, alt ca 2000 m, 26 Dee 1947, Carter, Alexander & Kellogg 2378 (DS); In granitic rocks at edge of Arroyo del Leén, N of road from La Paz to Los Planes, 22 mi SE of La Paz, alt ca 1565 ft, 11 Dee 1959, Wiggins, Carter & Ernst 494 (DS). This is one of the most widely ranging species among the species of Selayinella native to western North America, growing well through nearly 15 degrees of latitude, but mostly on the coastal slopes of the foothills and mountains within its range. It is also remarkable for the lack of variation between and within popula- tions, and for the diversity of the substrata it occupies. It seems to be equally adapted to granitic rocks, to basaltic extrusions, on sandstone, in crevices in serpentine, or gravelly detritus derived from any or all of these basic sources, and even on clay soil where little rock is present. It occurs too, on both north- and south- facing rocky areas if it is within the influence of coastal fog drift, and does well on vertical cliffs, gentle slopes or nearly level areas in shallow depressions. 4, SELAGINELLA CINERASCENS A. A. Eaton, Fern Bull. 7: 33. 1899. Forming close coverings of clay soil in open areas or among shrubs in coastal areas near San Diego and southward into Baja California. AMERICAN FERN JOURNAI. VoLuME 61, PLaTE 24 SELAGINELLA IN BaJA CALIFORNIA 157 SPECIMENS EXAMINED: Baja California: Among chaparral about 20 mi N of Ensenada, along road to Tijuana, 28 Feb 1934, Ferris 8474 (DS); Ensenada, mostly on hillsides, 7 Apr 1921, J. M. Johnston 3004 (CAS); about 30 mi S of Tijuana, 13 Mar 1956, Howell 30987 (CAS); Hillside NE of town of Ensenada, alt 200-400 ft, 23 Feb 1930, Wiggins 4213 (DS); about 12 mi N of Santo Tomas, along highway, 29 June 1962, Wiggins & Thomas 416 (DS). The paucity of specimens of this species in herbaria is doubtless attributable to the difficulty one has in freeing good specimens from the heavy soil on which it grows. When the substrate is moist it requires tedious, time-consuming effort to get the plants separ- ated from the clay, and when dry the earth is so hard that the fragile, closely attached plants are badly broken in the process. Consequently, most collectors ignore the close mats of moss-like growth. The wide differences in the sizes of leaves found on a plant, as pointed out by Tryon, is not applicable to the leaves taken on the upper and lower surfaces of the stem or branches at the same distance from the apex (or base) of a stem or branch, as indicated by Plate 24, figs. a-c. 5, SELAGINELLA EREMOPHILA Maxon, Smiths. Mise. Coll. 72(5) : 3, pl. 2. 1920. On shaded sides of rocks in open desert, along ledges and on steep to vertical walls of canyons, and occasionally on gravel at Figs. a-g, SELAGINELLA CINERASCENS: & = LEAF FROM UNDER SIDE OF E PROSTRATE STEM; b, ¢ = RESENTATIVE LEAVES FROM UPPER SIDE OF STEM; d = MicrosPOROPHYLL WITH MICROSPORANGIUM INTACT; e = MEGASPORO- HYLL, ABAXIAL FACE; f = MEGASPORE, CO URAL VIEW; g = M OUTER FACE; i = LEAF FROM UPPER SIDE OF DECUMBENT STEM, TERMINAL SETA DETACHED; k = LEAF FROM UNDER SIDE OF STEM, OPPOSITE i, TERMINAL SETA DETACHED; 1 = MEGASPOROPHYLL, ABAXIAL VIEW. Fics. m-t, SELAGINELLA LEPIDOPHYLLA: m = TIP OF BRANCHLET VIEWED FROM UPPER SIDE; D = SEG- MENT OF SAME BRANCHLET VIEWED FROM UNDER SIDE; 0 = LEAF FROM UPPER SIDE OF BRANCHLET, AXIS OF BRANCH TO ITS RIGHT; p = LEAF FROM UNDER SIDE OF BRANCHLET OPPOSITE 0, AXIS OF BRANCH TO ITS LEFT; q = MICRO- SPOROPHYLL, ABAXIAL VIEW; fr = SMALL STROBILUS AT TIP OF BRANCHLET, VIEWED FROM UPPER SIDE OF DORSIVENTRAL BRANCH; S = Micnosroun, LATERAL VIEW; t = MEGASPORE, OUTER FACE. 158 AMERICAN FERN JOURNAL the bases of such rocky areas, mostly below 3,000 feet, western Colorado Desert in California, in the Chuckwalla Mountains, ri v. Figs. a-~c, SELAGINELLA EREMOPHILA: a = NEARLY FULLY DEVELOPED LEAF FROM NEAR APEX OF BRANCHLET; b, ¢ = SUCCESSIVELY YOUNGER LEAVES FROM SAME BRANCHLET, ALL LEAVES STILL BEARING THE FRAGILE, TORTUOUS, : CELL e = LEAF FROM UPPER SIDE OF BRANCHLET; f = MATURE LEAF FROM UNDER SIDE OF PROSTRATE BRANCHLET, ABAXIAL VIEW; £ >= CELLULAR STRUCTURE OF APEX OF VENTRAL LEAF, ADAXIAL VIEW. California, and vicinity of Tinajas Altas, southwestern Arizona and south in Baja California about to the vicinity of Bahia de los geles. SPECIMENS EXAMINED: Baja California: Near Cantu Grade, 39 mi W of Mexicali, along highway to Alaska and Tijuana, alt ca 2500 ft, 6 Apr 1954, SELAGINELLA IN Baga CALIFORNIA 159 Wiggins 13045 (DS); Rocky hillside 0.5 mi SW of Rosarito, ca 40 mi SE of San Quintin, SW foothills of Sierra San Pedro Martir, alt ca 2900 ft, 16 May 1944, Wiggins 9981 (DS); Lower half of Cafion Diablo, east face of Sierra San Pedro Martir, near Lat. 31° 2’ N, Long. 115° 25’ W, 22-27 Mar 1954, Stombler 131 (DS); El Terminal, 16 mi S of Los Angeles Bay, common on N side of rocks near canyon mouth, near Lat. 28° 45’ N, Long. 113° 35’ W, alt ca 400 m, 14 Jan 1962, Moran 8524 (DS); About 6 mi S of Las Flores, S of Los Angeles Bay, near Lat. 28° 43’ N, Long 113° 32’ W, 13 Feb 1962, Wiggins & Thomas 272 (DS). The terminal seta on vegetative leaves of this species is re- markably slender and tortuous. However, this seta is very easily detached, so one rarely finds a leaf bearing a terminal seta except among the incurled, partially expanded young leaves at the very tip of a branchlet. Careful examination of a thoroughly moistened, relaxed branch apex under high magnification (about 30 to 40 X) usually reveals a few young leaves retaining the hairlike, white eta 6. SELAGINELLA LEPIDOPHYLLA (Hook. & Grev.) Spring in Mart. Fl. Bras. 1(2): 126. 1840. Lycopodium lepidophyllum Hook. & Grev. Bot. Mise. 3: 106. 1832. On limestone ledges, on talus slopes, in crevices in both cal- careous and igneous rocks, on north-facing slopes and shaded sides of boulders at contact between gravelly soil and the rock, from western Texas and New Mexico south through Mexico to El Salvador. SPECIMENS EXAMINED: Baja California: Peninsular divide W of Caduafio, Sierra de la Victoria, Cape Region, alt ca 4000 ft, 11 May 1959, Thomas 7859 (DS); On north-facing cliff overlooking river at Las Animas Ranch, near base of El Picacho, Sierra de la Laguna, alt ca 2000 ft, 12 Oct 1941, Hammerly 281 (CAS, Ds). The plants of both this and the following species often are called ‘‘Resurrection Plant” owing to the rapidity with which they flatten out and become bright green after being soaked by rain, in striking contrast to the apparently dead, compactly incurled branches during a period of drought. They can survive for months in the dormant condition, with the whole plant rolled into a grayish or brownish ball. The plants of both species are attached to the soil or rock only at the center of the rosette; no rhizophores 160 AMERICAN FERN JOURNAL appear along the under side of the flattened branches during periods of active growth and photosynthesis. Dead leaves and branchlets from a plant itself, vegetative debris from adjacent plants, dust, and soil accumulate around the bases and within the curled up balls of dry ones. This detritus often complicates the task of making acceptable herbarium specimens. 7. SELAGINELLA PALLESCENS (Pres!) Spring in Mart. FI. Bras. 1(2): 132. 1840. Lycopodium pallescens Presl, Rel. Haenk. 1: 79. 1 Selaginella cuspidata (Link) Link, Fil. Sp. Hort. he Bot. Berol. Cultae 158. 1841. Around rocks, at bases of cliffs and rocky ledges, on gravelly soil of meadows and in open stands of shrubs, Sonora and Chi- huahua, southward in Mexico and Central America into northern South America. SPECIMENS EXAMINED: Baja California: Canyon at La Laguna, Sierra de la Laguna, E of Todos Santos, alt 1650 m, 25 Dee 1947, Carter, Alexander & Kellogg 2335 (DS) ; Shaded rock faces along trail to Laguna from W, especially near summit, Sierra de la Victoria, 20 Aug 1955, Chambers 935 (DS); Potrero de Almerta, Arroyo de Hines, E slopes of Sierra de la Victoria, inland from Caduafio, among rocks alt ca 3400 ft, 9-11 May 1959, Thomas 7816 (DS); fares meadow surrounded by oaks and Pitas cembroides, La Laguna, Sierra de la Laguna, alt ca 6000 ft, 16-18 May 1959, Thomas 7930 (DS). Selaginella pallescens is aptly named, for when curled up in the dry condition, the under sides of the branches and the leaves on that side are silvery white, whereas the compact ball of a dry plant of S. lepidophylla has a reddish brown to rusty tinge. In both species the leaves on the under side of the branches are consider- ably longer that those on the upper side, the lower leaves extending appreciably beyond the outer margins of the upper ones. The stoutish, denticulate-scabrous seta at the apex of each leaf in S. pallescens contrasts sharply with the merely acute leaf of S. lepidophylla, which lacks a seta. This character can be seen easily with a hand lens, or in good light with the naked eye. Thus the two are not at all likely to be confused with one another. Dupiey Herparium, Stanrorp University, STanrorp, CA 94305. FERNS OF DOMINICA AND St. VINCENT 161 Notes on the Ferns of Dominica ard St. Vincent, IT C. V. Morton AND Davip B. LELLINGER This paper extends and concludes our notes on Lesser Antillean ferns and is based principally on collections made under the Bredin-Archbold-Smithsonian Biological Survey of Dominica that was active from 1963 to 1969, and also on collections made by C. V. Morton on St. Vincent in 1947. Since the prior report (Mor- ton & Lellinger, 1967) a few more botanists have visited Dominica and further researches by ourselves and others have solved some additional problems of taxonomy and nomenclature. DrPLazium AMBIGUUM Raddi, Opusce. Sci. Bologna 3: 292. 1819. Diplazium radicans sensu Christ, Farnkr. 220. 1896, pro parte. This South American species is known to us in the Antilles only from St. Vincent, where it was collected in the valley of the north fork of the Cumberland River at 400-600 m altitude (Morton 5469) and in the upper valley of the Richmond River at 330-540 m altitude (Morton 6182). These specimens match an isotype of the species found “in sylvis opacis prope Mandiocam, Brasiliae,” Raddi (FI, Morton photos 15914-15919 US). Diplazium ambi- guum closely resembles D. striatum (L.) Presl, which is widely distributed in the Antilles. The former species is bipinnate- pinnatifid, at least in most of the lower pinnae, whereas the latter is strictly pinnate-pinnatifid. It also resembles D. expansum Willd., but lacks the usually abundant hairs on the lower surface of the costae and costules of that species and has entire, not ciliate-erose, indusia. Diplazium radicans (Swartz) Desv. is not a synonym of D. ambiguum, but is a name based on Asplenium radicans Swartz, 1806, non L., 1759. (The two other names cited by Desvaux on p. 281 of his ‘““Prodromus”’ are invalid, later homonyms in Asplentum) Under A. radicans Swartz, Desvaux cites ¢. 76 of Schkuhr’s “Kryptogamische Gewiichse,” a dubious species according to Christensen’s “Index Filicum,” plus Swartz’ original publication 1 Partial cost of publication borne by the Smithsonian Institution. 162 AMERICAN FERN JOURNAL of the species, which is based on one of his own specimens from Jamaica. Judging by Maxon’s photograph of this probable holo- type and his annotation of the specimen, it is a taxonomic synonym of Asplenium dayi Hieron., which is the valid name for this ex- tremely rare Jamaican plant. ELAPHOGLOSSUM PUSILLUM (Mett. ex Kuhn) C. Chr., Ind. Fil. 314. 1905. This species has heretofore been known only from the Greater Antilles and from Guadeloupe and Martinique. The first record for Dominica (Wilbur 8097) is from the forested slopes just below the summit and from dense thickets on the summit-of Morne Trois Pitons, at 4000-4400 ft altitude. It is recorded as a common epiphyte. ERIOSORUS HISPIDULUS var. DOMINICENSIS A. F. Tryon, Contr. Gray Herb. 200: 115. 1970. Chambers 2596, from the summit of Morne Trois Pitons, is the first collection of Eriosorus from Dominica. It was collected in a habitat typical for this genus: in a dense growth of pendent moss, on the side of a large, exposed rock at the lower margin of the most dwarfed pigmy forest, at about 4400 ft altitude. PITYROGRAMMA TARTAREA (Cav.) Maxon, Contr. U. 8. Nat. Herb. 172173. 1913. Proctor (1965, pp. 219-220) has argued that the correct name for this species is P. ebenea (L.) Proctor, based on Acrostichum ebeneum L. Sp. Pl. 1071. 1753. He lectotypifies this name on the basis of sheet 1245.14 in the Linnaean Herbarium, which represents a small, depauperate specimen of P. tartarea. We disagree, and consider A. ebeneum to be a synonym of A. calomelanos [= Pity- rogramma calomelanos (L.) Link]. This has been the usual dis- position of A. ebeneum. The name A. ebeneum originated in a pre-Linnaean dissertation oe by J. B. Heiligtag in Linnaeus’ ‘““Amoenitates Academi- ” It included several phrase-name synonyms based on material a Jamaica, Martinique, and Brazil, plus a description of a FERNS OF DOMINICA AND St. VINCENT 163 large specimen apparently agreeing with P. calomelanos (‘Stipes pedalis & bipedalis”) that Linnaeus omitted from the account of A. ebeneum in the “Species Plantarum.” In that work the proto- logue is restricted to the following elements: 1. A citation (Amoen. Acad. 1: 272. 1749) with the phrase name “‘Acrostichum fronde pinnata, pinnis sessilibus oblongis sinuatis, summis brevissimis integerrimis’’ copied exactly from the ‘“‘Amoenitates.”’ Of the phrase-name synonyms in the “Amoeni- tates,” all but the two citations of Sloane’s names, which are the only ones based on Jamaican material, should be disregarded for reasons set forth below in point 4. Sloane’s names, descriptions, and plate are referable to P. calomelanos. The plate depicts a small, rather depauperate specimen. On the other hand, those Sloane descriptions and plate that truly do refer to P. tartarea (Voy. Jam. 35, t. 7, f. 1. 1707 and Cat. Jam. 21. 1696) are not mentioned by Linnaeus either in the “‘Amoenitates” or in the ‘‘Species Plantarum.” 2. A direct reference to Sloane, Cat. Jam. 20. 1696 and to Voy. Jam. 92, t. 53, f. 1. 1707, which are the two ice references in the ‘‘Amoenitates”’ mentioned above in point 3. A direct reference to Ray, Hist. Pl. ee 83. 1704, which is based on Sloane, Cat. Jam. 20. 1696, and thus also refers to P. calomelanos. 4. “Habitat in Jamaicae sepibus humidiusculis,” a statement of locality that restricts type material—specimens or literature— to Jamaica. The specimen in the Linnaean herbarium (1245.14) may or may not be a part of the protologue. Although it is anno- tated ‘‘Acrosticum [sic] 20. ebeneum”’ in the hand of Linnaeus, there is no evidence that it is from Jamaica or was in Linnaeus’ possession when either the ‘‘Amoenitates” or the “Species Planta- rum”’ was written. The phrase name in the ‘‘Amoenitates” that Linnaeus copied in the ‘Species Plantarum” does seem to apply to P. tartarea better than to P. calomelanos (“‘pinnis sessilibus oblongis sinuatis”’), and it could apply to the specimen in the Linnaean herbarium. But the more ample description in the ‘‘Amoenitates” that Lin- 164 AMERICAN FERN JOURNAL naeus omitted in the ‘Species Plantarum’ clearly does not refer to the specimen now in the Linnaean herbarium. The fact of this omission and the restriction of the type to Jamaican material indicates that si as concept is based primarily on ¢. 53, f. 1 of Sloane’s ‘‘Voya Tryon (1962, p. 60) suggested that Jenman (1886, p. 38) in- ferentially lectotypified A. ebeneum on the Sloane plate. However, Jenman’s paper is merely an attempt to identify the Jamaican ferns in Sloane’s herbarium; there are no synonymies in taxonomic form. Under Gymnogramme calomelanos Jenman places the Sloane phrase name and reference (Hist. Jam. 92, ¢. 53, f. 1. 1707), and adds ‘“‘Acrostichum ebeneum Linn.—a young plant [of G. calo- melanos|.’’ This, we believe, was Linnaeus’ actual concept. PTERIS ARBOREA L. Sp. Pl. 1073. 1753. Hodge (1954, p. 85) lists this species only for the islands adja- cent to Dominica, but it has been collected several times on the island at 1000-1800 m altitude (Hodge 3236 from near Salybia, Chambers 2535 and Lellinger 393 from near Pont Cassé, Lellinger 443 from near Trafalgar Falls, and Hodge 74 from Lisdara). The pedate and areolate species of West Indian Pieris related to P. altissima Poir. have long been confused, and many names have been in use for them. Specimens in the U. 8. National Herbarium can be separated easily with the following key: Fronds mostly 0.5-1 m long; costular areolae 2 (or 3 in large specimens), gi parallel to yee costa; costular spinules pointed and usually spread- BIDID AG LTAN-PTOCN is 5 oie ok ee bh es Se P. altissima Poir. Fronds aoa 1-2.5 m me: costular areolae 1 or 2, the distal one, if present, always parallel to the costule, not the costa; costular spinules absent, rudimentary, or weakly developed and usually parallel to the costae; Weignen Wight ee a a L. For the most part, we have seen type fragments, ‘souchentic specimens, or sometimes photographs of type specimens of the names we place in synonymy. Under P. altissima Poir. we include P. brevinervis (Fée) Jenm., P. bulbifera Jenm., P. elata Agardh, and P. kunzeana Agardh. Under P. arborea L. we include P. aculeata Swartz, P. crassipes Agardh, P. lactuca Poir. in Lam., P. longi- brachiata Agardh, P. multiserialis Jenm., and P. regia Jenm. FERNS OF DOMINICA AND St. VINCENT 165 The basis of P. arborea should be t. 5 of Plumier’s ‘‘Tractatus de Filicibus Americanis.” (In this same work t. 1/7 is sterile material of the same species, but was not cited by Linnaeus.) The other basic reference cited by Linnaeus, ¢. 3 of Plumier’s “Description de Plantes de l’Amérique,” is also part of the protologue of Polypodium spinosum 1L., presumably through a typographical error. Pteris hartiana Jenm. is a taxonomically distinct species from Trinidad. It has at least three areolae along the costae, one long and the others shorter; as in P. altissima, all are parallel to the costa. It differs from P. altissima in having 4-6, rather than 2 or 3, rows of areolae between the costae (or costules) and the margin. LITERATURE CITED Honae, W. H. 1954. Flora of Dominica, B. W. I. Part. I. Lloydia 17: 1-238. JeNMAN, G. 8. 1886. On the Jamaica ferns of Sloane’s herbarium. J. Bot. Brit. & For. 24: 33-48. Morton, C. V. and D. B. LELLINGER. 1967. ae on the ferns of Dominica and St. Vincent. Amer. Fern J. 57: Ti. oa G. R. 1965. Taxonomic notes on oo , ferns. Brit. Fern Gaz. 9: 213-221. Tryon, R. M., Jr. 1962. Taxonomic fern notes. II. Contr. Gray Herb. 185: 52-76. U. S. NavronaL Herparium, SMITHSONIAN INSTITUTION, WasuHineton, DC 20560. 166 AMERICAN FERN JOURNAL Two New Tree Ferns from Costa Rica Luts Dirco G6MEz The tree fern flora of Costa Rica is richly represented by some 50 or 55 species of the Cyatheaceae and Dicksoniaceae. Two have been known from Cocos Island (5°30’N, 87°03’W), a Costa Rican possession in the Pacific Ocean some 300 miles southwest of the Punta Burica Peninsula, the southernmost part of Costa Rica. These species, Cyathea notabilis Domin and Trichipteris nestotica (Maxon) Tryon (formerly known as Alsophila notabilis Maxon and A. nestotica Maxon), occur on the island from sea- level to its highest point, Monte Yglesias, about 900 m altitude. During an cight-week survey of the island’s cryptogamic flora, I noticed a tree fern that differed in several respects from the two previously reported species. Further study lead to the conclusion that this is a new species: vCyaTuHEa alfonsiana Gémez, sp. nov. nu ; inframediales, indusiati. Indusium squamiforme, parvum. “Tyre: Twin Mountains, Upper Wafer Valley, Cocos Island, ) AMERICAN FERN JOURNAL VotuME 61, PLate 25 oe a Say TR NACHT. ; MUSEO NACIONAL DE COSTA MCA Ho.orypr or Cy, THEA ALFONSIANA (CR). Fic. 1. Mepiau PINNAE. Fic. 2. MEDIAL PINNULES. SCALE IN CM. 168 AMERICAN FERN JOURNAL PARATYPES (identity confirmed by R. M. Tryon, Jr.): Cocos Island, Holdridge 5160 (GH), W. L. Schmitt 129, 130, 131 (all US). According to the recent classification of the Cyatheaceae by Tryon (1970), this species falls in Cyathea, mainly on the structure of the scales of the trunk, stipe, and rachis; that is, the scales are marginate with a broad margin of cells unlike those of the central portion. The stipe scales lack setae and have, like the rest, a filamentous apex. Various types of indument are present on the stipes, rachises, costae, and costules. The venation is free and there is a scale-like indusium present. The new species differs from C. notabilis and from T. nesiotica in general habit, but especially in having a scale-like indusium; the indusium is absent in T'richipteris and is sphaeropteroid in C. notabilis. Such peculiarities of the in- dusia of these Cocos Island tree ferns were first noted by Maxon in the specimens collected by Schmitt (C. V. Morton, pers. comm.). Cyathea alfonsiana is named after Mr. Alfonso Jiménez M.., former Curator of the Costa Rican National Herbarium, as a humble tribute to his many contributions to the general knowledge of his country’s flora. The second new species is an endemic fern of the central north- eastern slopes of the Talamanca Ridge in continental Costa Rica. This species is in evident danger of extinction due to uncontrolled deforestation. It presents a number of interesting characters, one of them of ecological interest. In collecting some immature sori for chromosome counts, I discovered that a moth of the Nymphalid group deposits its eggs in rows that duplicate exactly the disposition and appearance of the immature sori. All attempts to rear the larvae and adults of this remarkable insect failed, and the moth still remains unidentified. The new species is a conspic- uous, tripinnate tree fern also belonging to Cyathea in Tryon’s classification. It was formerly described, but unfortunately not validly or effectively published, by Nisman (1965) as Hemitelia holdridgei, and is here published as: A Cyatuea holdridgeana Nisman & Gémez, sp. no ncus rectus ca 2 m longus, inermis, ‘paleac se paleis stram- ineis ovato-lanceolatis, semiuncialibus, brunneis guttatis. Frons AMERICAN FERN JOURNAL VouLuME 61, PLATE 26 i. RRR Nard GAN SERRE eosin eke Sani MUNG NACCHAL SE CosTA WER MEDIAL PINNA. Houotyre or CyATHEA HOLDRIDGEANA (CR). Fic. 3. Fig. 4. MepIAL PINNULE AND PART OF RACHIS. SCALE IN CM. 170 AMERICAN FERN JOURNAL m longa, eg ios Stipes et rhachis semiteretes canaliculati, re opaci, parce muricati. Stipes 7 cm longus, 1.5 em crassus ad basim pa eaceus, eee furfuraceus, zp glabrescens; rhachis supra strigosa, subtus furfuracea, paucis paleis sparsis, pinnis 17- jugatis. Pinnae petiolatae (petiolo 2. 5 as longo), rhachibus fere teretibus canaliculatis, dense strigosis, subtus iat ORE squamu- losis, deltoideo-lanceolatis, pinnulis 20-jugatis; laminae apice acuminatae serratae. Pinnulae petiolulatae (4 mm) deltoideo- parte pinnulae distincta, sessilia, ad basim laeve inaequalia. Pars apicalis pinnulae pinnatisecta fere ad costam. Laminae coriaceae, supra glabrae, virides, subtus acy oath margine revoluto serrato. Costa secundaria ad basim strigosa, ad apicem cum 2-4 pilis rigidis, subtus squamulosa. Sori inframediales, orbiculares, trans- lucenti-virides, costis secundariis propin Type: La Chonta, at Km 55 of the ee Highway, Prov. Cartago, Costa Rica, alt 2200 m, Carmen Nisman S. 104 (CR; isotypes F, GH). ParatypeEs: La Chonta, Prov. San José, Costa Rica, alt 2300 m, Gomez 2542, 2577, 2560, Goldgewicht & Gémez 2675 (all Herb. Gémez); Km 54 of the Interamerican Highway, Costa Rica, J. A. Saenz (Herb. Gémez). Although there are a number of species of Cyathea with tri- pinnate or tripinnate-pinnatifid fronds, such as C. hemiotis Christ and C’. hastulata Christ, the closer affinity of C. holdridgeana seems to be with C. multiflora J. E. Smith and C. costaricensis (Mett. ex Kuhn) Domin, from which it differs in having sessile pinnae, petiolulate pinnules, a strictly inframedial sorus, and bicolorous and corticinate scales on the stipe. LITERATURE CITED Nisan S., Carmen. 1965. Estudio taxonédmico y ecolégico de los helechos arborescentes (Cyatheaceae y Dicksoniaceae) de Costa Rica. Lic. hesis, University of Costa Rica Tryon, R. M., Jr. 1970. The nia a of the Cyatheaceae. Contr. Gray He aes 200: 3-53. Museo Nacrionat pe Costa Rica, APaRTADO 749, San Jos#, Costa Rica. SPORE GERMINATION AND PROTHALLIAL GROWTH 171 Comparative Habitat Requirements for Spore Germination and Prothallial Growth of Three Ferns in Southeastern Michigan Royce H. Hi! Investigation of the physiological ecology of various plants has yielded information concerning mechanisms and patterns of evolutionary divergence (Bjérkman & Holmgren, 1963; Ayodele Cole, 1967; Hadley & Levin, 1969; Mooney & Billings, 1961). Such studies have largely omitted pteridophytes. Owing to the alternation-of-generations life cycle, certain problems arise con- cerning physiological divergence within groups of pteridophytes. The success of pteridophytes in a community is obviously limited both by the successful establishment of free-living gametophytes and the survival of sporophytes. Thus, the habitat requirements for spore germination and prothallial growth are critical factors for the success of the plant. Physiological divergence between populations or species, as a result of selective pressures of eco- logically different habitats, should be apparent in both phases of the pteridophyte life cycle. Farrar (1967) found fern gametophytes reproducing inde- pendently in the southern Appalachians, far north of the normal ranges of their sporophytes. Conway (1953) stated that there are few records of finding sporelings of Pteridium aquilinum in Scot- land, even though the sporophytes are common and spore pro- duction is heavy. She suggested that spores are released late during the growing season and that Pteridium sporelings are more susceptible to frost than are those of other species observed. Conway also noted that sporelings are subject to attack by soil insects and fungi. These two studies suggest that the gameto- phytic and sporophytic phases of a plant can have different habitat requirements and that one phase of the life cycle can be Dr. wee poeta as ey eG coke. 7 peg ere Botanical Gardens, University of Michigan. L72 AMERICAN FERN JOURNAL limiting to the success of the species in extending its geographical range and ecological amplitude. The purpose of this investigation is to compare the habitat requirements of prothallia of three polypodiaceous species that occupy ecologically contrasting habitats in southeastern Michigan. MetTHops Species from habitats differing in exposure to sunlight and in substrate pH were selected for study. The habitats are located in Washtenaw County, Michigan. Thelypteris palustris Schott (Marsh Fern) grows abundantly on the open sphagnum mat of Mudlake Bog. Woodwardia virginica (L.) J. E. Sm. (Virginia Chain Fern) occupies floating, graminoid mats in a swamp clearing at the Waterloo Recreation Area. Adiantum pedatum L. (Maidenhair Fern) grows in an upland northern woods at Sharon Hollow. Light intensity and temperatures were taken at the habitats. Light intensity was measured with a General Electric Type DP-9 light meter and temperatures were taken at 30 cm and at the sub- strate surface with glass mercury thermometers. Readings were taken at each habitat and at an adjacent, gravel road to obtain habitat and ambient conditions, respectively. The ambient readings were intended to facilitate comparisons of environmental data taken on different days, August 7 at Sharon Hollow and at Waterloo and on August 30 at Mudlake Bog. Both days were bright and hot, as indicated by the similarity of the ambient conditions of the habitats. These data appear in Table I. Spores were collected by washing fertile fronds under running tap water, blotting them dry, and placing them in envelopes for storage for several months at 6° C. Upon removing the spores from storage, they were filtered through a 0.177 mm sieve and several layers of cheesecloth to screen out extraneous material such as sporangia. They were then sown in 10 X 35 mm plastic petri dishes containing nutrient medium, at a density of approximately 100 spores per square cm. The medium was one of several varia- tions of full-strength Bold’s nutrient solution. Liquid medium and liquid solution added to blotting paper were usable, but these SPORE GERMINATION AND PROTHALLIAL GROWTH 173 techniques often led to fungal or algal contamination of the cul- tures. Bold’s solution solidified with 1% bacto-agar was the medium least susceptible to contamination. The petri dishes were placed under gradients of light intensity and pH, inside a growth chamber, and of temperature on a thermo- gradient plate. In the light intensity experiments, the light source was a combination of incandescent and Cool White fluorescent lamps; the petri dishes were placed at varying distances from the light source and/or were covered with layers of cheesecloth in order to vary the light intensity. Temperatures inside the plates were determined with a Telethermometer probe and maintained at 25 +1°C. Temperature control at the higher values of light intensity was facilitated by placing the dishes in the direct flow of cool air above the chamber vents and by positioning a plexiglass water screen containing 10 cm of water between the dishes and the light source. The pH of the medium was adjusted to 6.7 by adding NaOH or HCl for both the light and temperature experiments. Petri dishes were placed under 500 ft-c at 25 + 1°C in the pH experiments and under incandescent light at 500 ft-c in the tem- perature experiments. After a growth period of 3 weeks, the percentage of germination and prothallial width (to 0.1 mm) were determined, the former based upon 400 spores or prothallia and the latter upon the maxi- mum width of 40 prothallia, measured with an ocular micrometer mounted in a dissecting microscope at 40 X. Prothallial width was chosen as an index of growth because width increments of gametophytes appeared to be more directly related to increments of prothallial area than were length increments. Dark-grown prothallia in the light intensity experiments, for example, were long and spindly, whereas cordate prothallia at higher light in- tensities attained somewhat greater prothallial width but only slightly greater length than dark-grown prothallia. ENVIRONMENTAL DaTa Table I shows that light intensity (LI) was highest in the open habitats. The bog habitat is fully exposed to light except for 174 AMERICAN FERN JOURNAL occasional shade cast by poison sumac shrubs; the swamp habitat is shaded only during the early morning hours by an upland oak- hickory woods just east of the swamp. LI at the wooded habitat on August 7 varied from 77 ft-c in deep shade to 1192 ft-c where light flecks penetrated the canopy, even though ambient LI (at the road) was 5700 ft-c. Table I also shows that the air and surface temperatures, relative to ambient temperatures, were also higher in the open habitats than in the wooded habitat. Table I shows that the open habitats are acidic, whereas the woodland habitat is circumneutral. The bog—a typical northern Tasie I. CoMPARISON OF THE HABITATS OF THRBE FERN SPECIES Adiantum Thelypteris Woodwardia Species pedatum palustris virginica Locality Sharon Hollow Mudlake Bog Waterloo Habitat wooded upland open bog swamp clearing Light Intensity (ft-c) ambient 5700 4556 5479 habitat 77-1192 4300 4552 Temperature (°C) ambient (30 em) 34.0 33.5 35.4 habitat 30 cm 27.4 31.8 32.0 surface 27.2 30.0 Substrate rich humus sphagnum mat graminoid mats Substrate pH 7.6-7.8 5.5 5.9 one—is dominated by sphagnum, larch, and ericaceous shrubs. The swamp substrate is acidic and highly anaerobic. The wood- land substrate consisted of leaf litter overlying a deciduous forest mull. CoMPARISON OF GERMINATION TimES AND GROWTH RaTES Thelypteris and Woodwardia spores required about 2 days for germination, whereas Adiantum spores germinated in 5 days. The first two species also had faster growth rates than did Adiantum during the 3-week period following germination. Prothallial width SPoRE GERMINATION AND PROTHALLIAL GROWTH 175 of Adiantum never exceeded 0.6 mm, whereas that of the other species reached 2.0 mm at the lower pH values. The shorter germination time and faster growth rates of the open-habitat species may constitute an adaptive advantage. It is likely that prothallia of these species would have faster growth rates and be more competitive than would Adiantum in an exposed habitat. TEMPERATURE EXPERIMENTS Petri dishes of each species were placed on a thermogradient plate to establish a 15-35°C temperature gradient. In addition, dishes were also placed in two cold rooms at 5 and 10°C. As seen in Fig. 1, none of the spores germinated at 5°C, although spores did germinate after they were moved to 25°C following their exposure to the cooler temperature. Other experiments showed that none of the spores germinated at 40°C and that the spores lost viability, for they did not germinate when they were moved to 25°C. Thelypteris and Woodwardia spores germinated at 35°C, but Adiantum spores did not. The temperature limits of germination for Thelypteris and W oodwardia are approximately 10-35°C, whereas those of Adzan- tum are about 10-30°C. The higher temperature tolerance of the open-habitat species may be considered adaptive, since surface temperatures of the bog and swamp probably reach 30-35°C regularly during the mid-hours of the day. Surface temperatures of the woodland habitat probably seldom reach 30°C. Spores landing on the sphagnum mat of the bog or on the floating mats of the swamp would certainly be exposed to more adverse temper- atures than those landing on the forest floor of the woodland habitat. Figure 1 also shows that optimum or near-optimum tempera- tures for prothallial growth are higher for the open-habitat species than for Adiantum. These were 25-30°C for Thelypteris, 25°C for Woodwardia, and 15-25°C for Adiantum. Prothallial growth of the three species followed much the same pattern as germination, although it can be seen that the temperature limits of germina- tion are broader than those of maximum or near-maximum growth of prothallia. Although spores of Thelypteris and Woodwardia 176 AMERICAN FERN JOURNAL germinated at 10°C, prothallial growth was relatively low at temperatures below 25°C. By contrast, Adzantum prothallia had relatively high growth rates at 15°C. The preference for lower temperatures in the latter species, as compared to the open- habitat species, appears to be an adaptation to the cooler wood- land habitat. Growth of Adiantum prothallia, compared to the other species, was decidedly slower and dropped abruptly at temperatures above 25°C. The preference for higher temperatures Thelypteris Adiantum Woodwardia -—-- ante ee 100 gma Lebeau » ° ma “f <7) \ spr ah § 12 \ \ 7 N. FOO = on 5 \ - ae 460 E (a Sf a o = fa = B ‘ ft B20 z 5 \ x : A ff }# i i = = i i i = i a 10 615 «20.0: «(OSS 10-18 20 26 25 30 36 10 15 20 25 30 35 Temperature (C ) mean 1 standard deviation range ® %-germination 1G. 1. THE EFFECT OF TEMPERATURE ON PROTHALLIAL WIDTH AND PERCENT GERMINATION OF SPORES. and the tolerance of higher temperatures in Thelypteris and W oodwardia appear adaptive to the open habitats of these species. Thus the temperature requirements for prothallial growth of these species are even more in accord with their respective habitat condi- tions than are their temperature requirements for germination. The temperature limits of spore germination may be compared with those reported by Hartt (1925) for Onoclea sensibilis, in which the spores germinated at 16-34°C with an optimum of 28°C. Spore viability was lost at 40°C. Hevly (1963), in a study of the physiological ecology of cheilanthoid ferns, reported much higher SPorE GERMINATION AND PROTHALLIAL GROWTH 177 temperature limits. Spores of Notholaena cochisensis, a xerophytic fern that typically grows fully exposed to the sun, germinated at 10-50°C with an optimum of 25-35°C, whereas Cheilanthes fend- leri, which usually inhabits crevices around the bases of boulders, germinated at 0-35°C with an optimum of 15-20°C. The spores of the species investigated here germinate in only a few days after sowing, and previous experiments showed that freshly collected spores are fully capable of germinating. These observations suggest that spores would germinate in the field soon after their discharge from sporangia, and that the overwintering stage is the prothallium rather than the spore. Observations of seasonal phenology of these species showed that initial dehiscence of sporangia occurred on June 22 for Woodwardia, August 7 for Adiantum, and August 25 for Thelypteris. It would seem reasonable that spores shed late in the growing season, such as those of Adiantum and Thelypteris, would over- winter before germinating. If this were the case, the spores would probably have a chill requirement for germination. This was not found to be the case for these species, but the question of whether spores that fail to germinate during the growing season in which they were discharged could survive winter conditions arises. It was noted that the spores retained their viability after several weeks of storage at 6°C and at shorter exposure to 5°C on agar medium. In an attempt to resolve this question, spores were sown in a liquid medium which was then frozen for 30 days and then brought to room temperature. A high percentage of the spores germinated, and in the time periods characteristic for each species. It appears that spores of these species are able to retain viability under winter conditions, although the evidence suggests that the usual overwintering stage is the gametophyte rather than the dormant spore. These are only tentative observations and need to be followed by intensive field studies. Licgut IntTENSITY EXPERIMENTS A LI gradient of 100-3000 ft-c was established. In addition, petri dishes were covered with black cloth, after receiving an 178 AMERICAN FERN JOURNAL initial light stimulus during sowing, to effect dark conditions. Germination percentages and prothallial growth for the species over the gradient are shown in F%g. 2. Although spores of all species germinated at all LI values, the percentage of germination was lower in darkness and high germi- nation rates were reached at 100-3000 ft-c for all three species. Thelypteris Adiantum Woodwardia 7. hee =e Oe ed on - _——. —e | | bre % — germination prothallial width (mm) @o : i ft. eae x Ea 3 6 7 _ 7 A Serene: 5 7 Ls 7. v 7 —— Le | ee “Re Bee ln ease cote | | Sea rae eek Pe pee Light Intensity (10%, f-c ) * mean * 1 standard deviation I range * %—germination Fic. 2. THE EFFECT OF LIGHT INTENSITY ON PROTHALLIAL WIDTH AND PER- CENT GERMINATION OF SPORES The prothallia of all three species were long and spindly when grown in darkness. The prothallial width of Thelypteris was greatest at 1000 ft-c, and that of Woodwardia and Adiantum at 500 ft-c, but prothallial growth dropped off more abruptly at values above 500 ft-c in Adiantum than in the open-habitat species. The relatively faster growth rates of the open-habitat species, as compared to those of pape siete their selective advantage in establishment in open com The apparent chlorophyll nae e ie also indicate that the open-habitat species have higher tolerance to high LI than does Adiantum. Spores of all three species germinated at 3000 ft-c, SPORE GERMINATION AND PROTHALLIAL GROWTH 179 but the resulting prothallia were chlorotic as well as small. At 2000 ft-c, prothallia of Thelypteris and Woodwardia were pale green, whereas those of Adiantum were devoid of green color. Darker green colors were found in all three species at 100-500 ft-e. Spores of all three species germinated in darkness after an exposure of only a few seconds to the fluorescent light used during sowing. Mohr (1963) stated that light is an almost universal re- quirement of spore germination and prothallial development among ferns. Borthwick et al. (1952) noted that certain small seeds require a dose of red light for germination, a stimulus that can be nullified by a subsequent treatment with a far-red light. To determine whether spores of the species under study have no light requirement for germination, or whether the brief exposure of the spores during sowing to fluorescent light (including red light) satisfied a possible light requirement, spores were sown under neutral light only (absence of red light) and then placed in darkness After a period of two weeks, no spores had germinated, indicating a red-light requirement of these spores. It is likely that the germi- nation of these species is under operation of a red and far-red reversible photoreaction. Du Buy and Neurnbergk (1938) state that Pleridium aquilinum spores germinate well in light of “high intensity.” This is in accord with results reported here for Woodwardia and Thelypteris, but Hevly (1963) found that desert ferns of the Southwest germinated at a somewhat lower maximum LI. Miller’ and Miller (1961) found that prothallia of Onoclea sensibilis grew much more rapidly under 400 ft-c than under 220 or 28 ft-c. It was also observed in the present study that prothallia of all three taxa grew faster at 500 ft-c than at lower LI values. PH EXPERIMENTS Spores were placed in a pH gradient of 4-10 (Fig. 3). Thelypteris and Woodwardia, which grow in acid habitats, showed lower pH requirements than did Adiantum, whose woodland habitat is circumneutral. Spore germination of Thelypteris was high over the entire gradient, but prothallial growth showed a marked pref- 180 AMERICAN FERN JOURNAL erence for pH 5 and decreased sharply at higher pH values. Assuming that the germination process is adapted to the natural habitat of this species, this is in agreement with the observed pH value of 5.5 for water squeezed from sphagnum of Mudlake Bog. W oodwardia, whose habitat substrate pH was determined to be 5.9, had both the highest percentage of germination and the Pas ata ae -—-. son -e-- “100 \ / 455 ate ‘ » / \ / oe bo cs Thelypteris Adiantum Woodwardia p40 : ; al =} = & ey odt a | i ae i i: } . t i uae TG SES Coes em Weed See uae an. en lew eee 10 mean @ 1 standard deviation [range " %-germination Fic. 3. THE EFFECT OF PH ON PROTHALLIAL WIDTH AND PERCENT GERMINA- TION OF SPORES. fastest growth rates at pH 5-7. Percent-germination for Adiantum, whose soil pH was 7.6-7.8 in several samples, was high at all pH values except pH 4, and prothallial growth was maximal at pH 8. Again these species differences appear to reflect habitat differences. Fig. 3 shows that maximum prothallial width of Thelypteris and Woodwardia exceeded 2.0 mm at low pH values, whereas prothallia of these species grown in the temperature and LI experiments were smaller. This is probably a result of the pH (6.7) used in the latter experiments, which is higher than the optimum SPporE GERMINATION AND PROTHALLIAL GROWTH 181 pH values for these species. The comparative growth rates of these three species are also very apparent in the pH experiments, as prothallia of Adiantum at optimum pH (8) reached a maximum of only 0.5 mm in diameter. This contrast dramatically demon- strates the faster inherent growth of the open-habitat species as compared to Adiantum. It has been accepted for some time that fern species have definite habitat requirements for substrate pH. Wherry (1920) listed pH ranges for several rock ferns, and Kruckeberg (1964) reported that certain fern species of the Pacific Northwest are restricted to localized ultramafic rocks. It is reasonable that gametophytes have pH requirements similar to those of the sporo- phytes. McMillan et al. (1968) attributed the narrow distribution of Pilularia americana in Texas partially to the inability of spores to germinate on limestone soils. Spore germination of this species was inhibited at pH values above 6.4, although sporelings trans- planted to limestone soils grew at normal rates. The habitat of Pilularia in central Texas has a pH of 5.2-6.1. Hevly (1963) found a correlation between the pH of habitats of certain cheilanthoid ferns and their corresponding pH re- quirements of germination. Notholaena cochisensis, which grows on limestone soils, germinates at pH 7-9 with an optimum of pH 8. Notholaena grayi germinates at pH 5-8 with an optimum of pH 6 and grows on soils on igneous rock. The different pH requirements of the open-habitat species and Adiantum, like the different temperature and LI requirements of these species, reflects the different habitat conditions of these species. These observations provide strong evidence that the processes of spore germination and prothallial growth of these species, as a result of variation and selection, have undergone physiological divergence. LITERATURE CITED AyYopELe Cote, N. H. 1967. Comparative physiological ecology of the genus Eriogonum in the Sats Monica Mountains, Southern California. Ecol. Monogr. 37: 1-2 182 AMERICAN FERN JOURNAL Bs6RKMAN, O. and P. Hotmeren. 1963. Adaptability of the photosynthetic apparatus to light intensity in ecotypes from exposed and shaded habitats. Physiol. Plant. 16: 889-914 BortTHWICK, H. A. ET AL. 1952. A reversible photoreaction controlling seed germination. Proc. Nat. Acad. Sci. 38: 662-666. Conway, Exsre. 1953. Spore and sporeling survival in Bracken (Pteridium aquilinum (L.) Kuhn). J. Ecol. 41: 289-294. Du Buy, ‘E G. and E. L. NeurnsperGk. 1938. Growth, tropisms, and other movements, Chapter X in Manual of Pteridology, ed. F. Verdoorn. M. Nijhoff, The Ha, Farrar, D. R. 1967. Gametophytes of four tropical fern genera reproducing independently of their sporophytes in the southern Appalachians. ience 155: 1266-1267. Haptey, E. B. and D. A. Levin. 1969. Physiological evidence of ‘eee! tion and ee evolution in Phlox maculata. Amer. J. Bot. 56: -570. Hartt, Constance E. 1925. Conditions for germination of spores of Onoclea sensibilis. Bot. Gaz. 79: 427-440, pl. XXIX. Hevty, R. H. 1963. Adaptations of cheilanthoid ferns to desert environments. J. ar Acad. Sei. 2: 164-175. KruckesBerG, A. R. 1964. Ferns associated with ultramafic rocks in the Pacific Northwest. Amer. Fern J. 54: 113-126. McMitian, C. Er at. 1968. Factors influencing the narrow restriction of Pilularia americana in Texas. Southw. Nat. 13: 117-127. Miter, J. H. and P. M. Mixer. 1961. The effect of different light conditions and sucrose on the growth and ip pomagns of the gametophyte of the fern, Onoclea sensibilis. Amer. J. Bot 154-159. Monr, H. 1963. The influence of visible artes on the germination of S—— spores and the growth of the fern protonema. J. Linn. ¢. London, Bot. 58: 287-296. Mooney, i A. and W. D. Brings. 1961. Comparative physiological ecology of arctic and alpine populations of Oxyria digyna. Ecol. Monogr. : 1-29. 31 Wuerry, E. 1920. The soil reactions of certain rock ferns. I, II. Amer. Fern J. : 15-22, 45-52. DEPARTMENT OF Botany, UNiversiry oF MicHIGAN, ANN Arsor, MI 48104 GAMETOPHYTE OF THELYPTERIS ERUBESCENS 183 The Gametophyte of Thelypteris erubescers LENETTE R. ATKINSON Thelypte.is erubescens, the Reddening Thelypteris, is a fern of the mountains with a thick, creeping rhizome and a leaf that can be 6 feet tall or more. The stipe and large veins may be purplish- tawny (Beddome, 1892, p. 289) or wine-red when dried (Ching, 1963, p. 320). It is one of a small group of ferns in North India and West China and has been known for some time. Iwatsuki (1965, p. 178) in his study of the thelypteroid ferns says of it, the “‘dis- tinctness of this large and handsome fern has been recognized by various authors since Wallich (1828) gave this name to his Nepalian specimen.”’ Ching (1963, p. 320), who had examined the Wallichian collection (see Ching, 1936, p. 294), says “It has been long noted that of all the Chinese thelypteroid ferns, the well-known ex- indusiate Dryopteris erubescens (Wall.) C. Chr. from the China- Himalayan region always appears to be foreign among all the known genera, in none of which can it possibly fit.’”” Wallich called it Polypodium; other authors, depending on which characters were thought to be more important, have placed it in Glaphyropteris, Phegopteris, Nephrodium, and Dryopteris. Those who today would place this fern in Christensen’s tropical American subgenus Glaphyropteris (on the basis of habit and aerophores present) would do so against Christensen’s (1913, p. 158) judgement. In discussing his new subgenus Glaphyropteris he says “I do not know any Old-World’s species which can safely be referred hereto. The Himalayan D. erubescens (Wall.) C. Chr. is similar in habit but otherwise quite different.’’ Ching (1963, p. 320) concurred and placed it in his newly created genus Glaphyropteridopsis, the type species of which is Thelypteris erubescens. The chromosome num- ber reported for this fern by Mitui (1968, Lastrea erubescens) is n = 72 (on the base 36, one of the more common of the various numbers reported for the Thelypteridaceae). The gametophyte also shows the general characteristics of the group. I am indebted to Dr. R. E. Holttum for spores of Thelypterts 184 AMERICAN FERN JOURNAL erubescens (Wall. ex Hook.) Ching from a plant from Nepal growing at Kew Gardens, England. Dr. Holttum writes me that this “Kew plant ... was much smaller than the size to which plants of erubescens commonly attain.’ He continues, “In the Kew her- barium are several quite small specimens, much like that from which your spores came and also some very much larger, with pinnae to 35 cm. long and 3 cm. wide. There are also some rather intermediate in size.’”’ The gametophytes were grown under fluores- cent light on Knop’s solution solidified with agar (Atkinson, 1970). The bilateral spores are brown singly, black in mass and measure about 33 X 21.5 yw, exclusive of the spore coat. They are orna- mented by more or less discrete little wings which when dried are sharp-pointed and give the spore a shaggy or spiny appearance. A colorless rhizoid emerges first at germination. A 4—6-celled filament is formed whose cells may be greatly elongated when developed from spores retained and germinating within the sporangium. The first division initiating the cell plate is usually oblique in the terminal cell of the filament; occasionally it is vertical and in the subterminal cell. A cordate thallus is formed by the activity of an apical cell which may be replaced by an apical meristem as early as a month after germination. The margins of a month old thallus are wavy and may bear 0-3 hairs. At two months simple hairs may also be found on the surface. The thallus bears sex organs by 3.5 months, has a broad cushion and large wings with irregular margins. At 6 months the wings are very large, with folds in the surface and shallowly notched margins. Regeneration in older thalli takes place from the margins of the wings, and little groups of cells project irregularly from the sides of the cushion on the ventral surface. These may bear glandular hairs and abortive antheridia, and are interpreted as regenerative outgrowths. The first hairs are simple, chlorophyllous and marginal. They are late in appearing; some cordate thalli at one month do not have any hairs. They become glandular and the secretion is yellow. At 2 months they appear on the surface near the notch but are not abundant in either position; one thallus with 2 surface hairs had none on the margin. GAMETOPHYTE OF THELYPTERIS ERUBESCENS 185 The rhizoids are colorless or palest tan, long but do not tangle, and are abundant on the cushion, hiding the older sex organs. When thalli overlap in culture, rhizoids are often borne over the lower part of the cushion on the dorsal surface. The sex organs are of the advanced type. Archegonia appear at 2 months and embryos at 3.5 months, although no effort was made to produce them. Antheridia are scarce, and those first seen were on separate, usually smaller thalli. At 6 months they appear also at the sides of the cushion beside the opened and younger archegonia. Dehiscence was not observed. The large, green thallus with simple, yellow glands, ruffled wings, abundant colorless or pale tan rhizoids, and sex organs of the advanced type is not out of place among the gametophytes of the Thelypteridaceae, but it may also be found elsewhere among the ferns. The spore is not very large (as is sometimes the case with polyploids), but, then, spore size at the species level may not always accompany an increase in chromosome number, as Bir (1966) has reported for some Himalayan ferns. Dr. Holttum sug- gests that a field study of Thelypteris erubescens is desirable in a place where the species grows abundantly. No notable gametophytic features are observed in Thelypteris erubescens to suggest relationships to other ferns within the family. LITERATURE CITED re LenetTeE R. 1970. The gametophyte of secepneage conjugata and reypteris irregularis. Phytomorphology 20 mecca R. H. 1892. Handbook of the ferns of British india: Ca leu Bir, 8. S. 1966. Consiaiing between spore-size and pla evel i in ihe Himalayan asplenioid and athyrioid ferns. J. Palynology 2: Cuina, R. C. 1936. A revision of the Chinese and Sikkim-Himalaya haa 2 with reference to ge species from neighboring regions. Bull. Fan Mem. Inst. Biol. 6: —347. . 1963. A ee als of the family Thelypteridaceae. Acta Phy totax. Sin. 8: 289-335. CHRISTENSEN, C. 1913. Monograph on pe genus Dryopteris I. KX. Danske Vidensk. Selsk. Skr. VII, 10: 55-282. Iwatsvux1, K. 1965. Taxonomy of the thelypteroid ferns with special reference to the species of Japan and adjacent regions. IV. Enumeration of 186 AMERICAN FERN JOURNAL the species of Japan and adjacent regions. Mem. Coll. Sci. Univ. Kyoto, B, 31: 125-197. Mirv1, K. 1968. Chromosomes and speciation in ferns. Sci. Rep. Tokyo Kyoiku Daigaku, B, 13: 285-333. AMHERST COLLEGE, AMHERST, MA 01002. Shorter Notes A REcENT Finp or Isoéres 1n LovuistANa.—A population of about two dozen plants of Isoétes melanopoda Gay & Dur. has been discovered in Caleasieu Parish, Louisiana. This is the first collection of [soétes in the state since the 1939 collections of Brown and Lenz, and only the fourth collection of the species in Louisiana. Brown and Correll! reported three sites, one each in Avoyelles, East Baton Rouge, and Rapides Parishes. The nearest to ours is in Rapides Parish, approximately 75 miles northeast of Lake Charles. Correll? reported J. melanopoda f. pallida (Engelm.) Fern. in Harris County, Texas, approximately 150 miles west of Lake Charles. The plants we found grew with sedges in the marginal zone of a depression in a meadow. The plants have sporophylls about 20-30 em long, and all the collected specimens were fertile. These speci- mens have pale leaf bases, rather than the dark brown to blackish brown leaf bases of the typical form. Voucher specimens (J. Brooks & R. Maples 817) have been deposited at McNeese State Univer- sity and the University of Southwestern Louisiana.—JAmEs H. Brooks AND Rosert 8. MAapues, Jr., McNeese State University, Lake Charles, LA 70601. 1 Brown, C. A. and D. CorrE.L. 1942. Ferns and Fern Allies of Louisiana. L.S.U. Press, Baton Rou 2 CORRELL, D. 5S. 1956. aa and Fern Allies of Texas. Texas Res. Founda- tion, Renner SHORTER NOTES 187 JAMESONIA ALSTONIT IN Oaxaca, Mexico.'—The excellent monograph by A. F. Tryon,? treating all of the species of Jamesonia, gives the northernmost station for the species J. alstonii and for the genus as Voledén Tacand, which is on the border between Guatemala and the state of Chiapas in Mexico. While on a bryo- logical foray into the Sierra Judrez, the authors encountered among the grasses and herbs under scattered trees of Pinus rudis a fern which was not immediately recognized. Eventually the senior author realized that it was a Jamesonia. He tentatively identified it as J. alstonii, an identification kindly confirmed by Dr. John T. Mickel of The New York Botanical Garden. The fern was collected in the highest gap on the road between Oaxaca city and Tuxtepec, Oaxaca, at ca 9000 ft altitude, on 26 December 1970 (Sharp & Webster 720A, NY, TENN) The occurrence of this fern at this station is of interest not only because it represents an extension of its range well into Mexico, but also because this is a locality from which bryophytes of geo- graphical significance have been collected. Gymnomitrium lace- ratum, originally described from Japan but also known from the Great Smoky Mountains, Peru, and southeastern Africa, was reported there by Sharp and Hattori.* Conostomum pentastichum was also reported from the same station by Bowers and Sharp; except for Mexico and Costa Rica, it seems to be restricted to the southern hemisphere. The senior author also has collected the rather rare Guamatela tuerckheimii Donn. Smith (Rosaceae), originally described from Guatemala, below this gap on the eastern slope. It seems that this area is of great significance in providing valuable phytogeographical data.—A. J. SHarp and H. J. Wessrer, Department of Botany, University of Tennessee, sorely, TN 37916 1 Contribution from the Botanical Laboratory, a of Tennessee, N.S. 365, and from the Itasca Biology Session, University of Minnesota. ? Tryon, Alice F. 962. A m one ah of the fon ae Jamesonia. Contr. 188 AMERICAN FERN JOURNAL Recent Fern Literature : A MonoGRaAPH OF THE FERN Genus Errosorvs, by Alice F. Tryon. Contr. Gray Herb. 200: 54-174. 1970.—Alice Tryon has produced a model monograph of Eriosorus, the result of many years of painstaking work. She has studied the cytology, the gametophytes, the anatomy and morphology in detail; she has studied the plants in the field in Costa Rica, Brazil, Puerto Rico, Columbia, and Peru; in addition she has visited Europe several times and has seen most of the historic types. The taxonomic work can therefore be considered definitive. Although there are a few novelties described the general result has been to reduce many of the described species to synonymy. The perhaps controversial part of the work is the recognition of several plants as hybrids with the genus Jamesonia. This is reasonable in the case of Eriosorus elongatus (Grev. & Hook.) Copel., which is considered a hybrid between EF. cheilanthoides and .Jamesonia. These plants do grow together in Andean pAramos, and E. elongatus does appear somewhat intermediate in morphology. The matter is not so clear in the hybrids indicated as E. hispidulus X Jamesonia, which seem to show little Jamesonia influence. Some other doubtful plants are mentioned as variants, and so it seems that even in the case of a definitive monograph there will always remain some unanswered questions.—C.V.\ Index to Volume 61 Acrostichum, 19, 41, 97-101, 119; aureum, Antrophyum, 41 97, 98, 101; calounelance. 162; ebeneum, An Appendageless Psilotum. Introduction to 162-164; fesulllen. 19; serrulatum, 117; Aerial wanage Morphology, 75 speciosum, 101 Aspidiaceae, 42 Actiniopteridaceae, 74 A spain azescens 29; blumei, 145, 148; Adiantum, 71, 90, 92, 174-181; capillus- germanii, ba mmatosorus, 148; mac- Seen 43, on *. 90; cuneatum, ~ peda- pe Se: 24; stipulare, 26; , 88, 89, 172-174; raddi 95 roe So 2 Alsop, "142, ne nesiotica, 166; nakabilie A pleniopsis, Asplenium, se adiantoides, 43, tripin- si American Species of Plagiogyria sect. natifidum, 44; dayi, 162; i ian cn, 140; Carinatae, 110 mtchatkanum, 140, 141; kamtchaticum, Anaxetum 141; pinnatifidum, 32-34, 45, 47; pinnati- Anemia ake 137 fidum X trichomanes, 32, 33; platyneuron, INDEX TO VOLUME 61 3; radic aa aes ; polyodon, 5, 47, 45, 88, 89; robustum, 44; se asa 32-34, Bie enium kamt pe mea rere a Dubi- ous Eastern Asiatic Fer Asplenium pinnatifidum <8 eh 32 N ew Record for Indiana, owas a filix-femina, 88, eH Var. 140, e Ga supe hia of s, , africana, 1, 7; rarer e 7; imbricata, i 9, var. pinnate, |7,;.9.-11, 12; pinnata, 1, 3, 7-9, 12, var. ae is var Cet ky 9, 12 Azollaceae, 42 Belvisia, ‘3; igs 43 Biechnaceae, 74, Blechnu 110; ¢ ear 42; diversifolium var. paleaceo-setosum, 44; irregulare, 44; urbanii, 118; vieillardii var. si ; Bolbitis, 17; adeest os, Botrychium disse f. obliquum, 45; Virgini , 45, Boutin, : pi ecg for Psilotum nudum in Sonora, ago pe aay 7 ‘ tans, ar A Recent Find o id in ei Brow = Flore de ia grieve eiee: endances, . 3, Ptéridodophytes a. ‘ ‘41 Bulbous Adder’s-tongue Common in Louisi- a, 39 hi ph llus, 45, 47, 88, 89 oa li ; ae se fendleri, 177; lanosa, 47, 88; setigera, Cibotium, 51, 55 Cnemidaria, 142, 143 Cc Habi Requirements tat for rmination and Prothallial Growth Spore Ge Three Ferns in Southeastern Michigan, Cteni hers an 143 ee see ipvane vc var, alfon sis, lata, 44; nsiana 170; hastulate, 170; ip poe falda geana, 168-170; multiflo 166, 168; propinqua, 43; subg. Sphaero teris, 43; vieillardii um f m, 44, 88, Soong Hea buen, a7 aie 45 vallia, 96; valdivi: 96 aude taedtia p piiactss sch 45 Dicksoniaceae, 1 Diplazi ora, 170; ani um, 147; acrostichoides, 45; ambigu- Dryoathyrium, 140; pterorachis, 1 x Cama | 189 um, 161; echinatum, 42 161; pyenocarpon, 45; ellhank tel: site, 61 Dryopteris, 124, 83; 19; campyloptera, 124, 126; campyloptera x » ket Isa 124, 17; mm 4; var. landslosa, 37: spin a A, var. interm — yarf Ostrich Fern, Elaphoglossum sabe 162 Equisetaceae, 74 quisetum ae male Eriosorus, 188; chellanthoides, 188; elongatus 18 id 188, v 8; his lus, minicensis, 162 Fern Collections in pee European Herbaria—V he Fern Vegetation of — Atoll, 97 Fosberg, F. The Fern Vegetation of Aldabra ae 9 Fox, ells, Schemochromic Wi Blue Test: guriacés, of Selaginella, 137 The Gametophyte of Thelypteris erubescens 183 The Gametophytes = — Hybrids in the Fern Gen ty fev Gastony, G. J eral pinnat > 4 _trichomanes~ A Ne v Record for tes 32 he Genus ese in oe California, ee The reat ae ena J, Smith Description of a New Species, 119 -A nhs arf Ostrich Fern, 35 with Gibson, H. Badric Glaphy ropteriopsis, 183 leiche 73 Sine oe Two New Tree Ferns from osta Ri Grammitidaceae, a mmatogsorus, 147, 148; blumeanus, 145, Gym calomelanos, 1 Hemitelis, 13; hold ne Heteron 7 oides Heusser, arte Je Po oi waa gaa of Chile (rev.), 95 190 Hill, R . Comparative Habit ments a Spore oe lial Growth of T t Require- ation and Prot chal- Michigan Hills, L. V. (see A. Sweet) Holttum, R. e Genus ee J, = ith with Description ofa w Species, till aki, Barbara Joe, The Genus Adiantum in Cultivation Pals sdiaasects (rev.), 95 Howell, Ferns an ern Allies ra nee oe Nevada ain Californiaand Neva re Hiymendlepi 43; revoluta var, planiuscula, Hymenophyilscene, 73, 74 phy m le-ratii, 43; mnioides, 44, een 186, f. pallida, 186 8 1 Lellinger, B. The American ay Sacra of Plagiogyria sect. anerrage bie Lellinger, B. (see C. V. Morton) pe ang — Lindsaea, 5 pe sensi 49, 51, 53-58, subsp. 8 8 ens leg 55; subg. Lind: = 3h aha! , 14, 16, var darjeelingensis, 14-16, r. odorata, 16, var. = 16; repens, 55, s Schizolom 5s (Schizoloma) ensifolia Swartz in Haw: Lindsteacene 42, 74 Lomaria adnata, 111; areolaris, 122; arguta, dos 1; “cenuifo ole, a 119 Lommasgon, CC. 2-6. Young, Ir: Vascularization of Fern Leaves, 87 hosoria, 143 Lunathyriu: 140; sect. Athyriopsis, 140; bory: num, 1 140; ioe, Dryoathyrium, 140; japonicum, 140; sect. Lunathyrium 0 1 pteror; is, Be Lycopodium, 137; lepidophyllum, 159; nutans, 43; rs ol bet agai 43; Ua sum va i m, Macrothelypterig Stree ns mo aples, R. S8., Jr. e J. H. Brooks) Marattia rolandi-principis, 43, var. sub- bipinnata, a Marattiac Marsilea, 101, opt 109; berteroi, 104, 107; fournieri, 104, 107, 108; mac ropoda, 104, 107; mexicana, 102, 103, 105, 107; minuta, AMERICAN FERN JOURNAL 108; mollis, 104, 107, 108; mucronata, y ncinata, 104, 108; vestita, 103, 104, 108; u 103, 8 Marsileaceae, 74 atteuccia, 37; struthiopteris, 36, var. pensylvanica, = ati ium, 17, 18; fendleri, 19; macrophyl- 17; ‘18; Pitas aie 19; sorbifolium, Metaxya, 143 Mickel, J.T. & F. V ont Leaf Epidermal tudies in Marsilea, 10 Microlepia, les speluncae on. . Asple enium 4 hatkanum aa a Dubi astern Asiatic Fern, 140; The Collections in Some Euro- pean Herbaria—V ; The Genus Gram- matosorus, 145; pera Disposition of Menis macro um Kunze, 17; A Wrongly Loc: Raat Sect s of Pyrrosia, 94 Morton, C. V. Lattin r, Niphidium longifolium, a Asis sary New Combina- Ne si allnaae 183; albescens, 26; kunthii,28; ophyllum, 17 idcaheecee 51, 55; aan 97; cordifolia, 43; exaltata, '51, 52, 88, 89; floccigera, . saad Loc alt sl for ete nudum , Mexico, 141 A New Variety of Lindsaea odorata Roxb. from Darjeeling in the Himalayas, 14 pe 37. ; americanum, 37, 39, ghiiven Sie Niphtum longifolium, a Necessary New mbination Niphobolis 94; americanus, 39; cuneatus, 94 e Ferns of Dominica and St. eo at ae 16 Notholaena Scchinicnadis: 177, 181; grayi, 181 eaule, 51, 57, 58 O; esas 7: Pellaea, 128, 129, 133; andromedaefolia, 135; atropurpurea, 88, 35; bridgesii, 128, 133; b esii X mucronata, 130, ; ne mucronata, 133; longimucronata X w 34; mucronata, 128, 133; raid ate 133; wrightiana Pessopteri hte ttee9 ris, 183; macrophylla, 17 a haa aureum, 88-90 ERRATA Pilularia, 181; ec esiapee 18 Pityrogramma melanos, a 163; ebenea, 3 censis, 113 ; denticulata, 116; euph- lebia, 111; subsect. Euphlebiae, 111; sect. lagiogyrae, 111; thoi, ¥/. DIG; latifolia, 116; maxonii, 113, 1 btusa, 114; pectinata, 113-115; ee 111 subsect. Pycnophyllae 1; semicordata tid, -Sis—1i5 117: pai ies 113, 118; urbanii, 118 pie C. C. A Manual of Plant Names (re Poeilopteri meniscioides, 17 Polypodiaceae, podi 183; americanum, 39; den- longifolium, 39; molle, 24; e 26; polypodioide es, 5 he metophy tes of Natural ybrids the Fern Genus Pellaea, 18 The el Disposition of Meniscium macro phyllum Kunze, 17 Pustaceas, 42, 74 89 Psi “ee 75-77, 81-85, ve complanatum $1, 82; nudum, 75, 82, 1 Pteridaceae, Pteridium, roe aquilinum, 171, 179, var. latiugsculum, 45 aculeata, 104; altissima Pteris, 119, 144; 1 164; lactuca, multiserialis, 164; regia, 164; vitt ata, 49, Pyrrosia, 37, 39, 94; sere 39; lnsifoia, 39; rupes estris , 94; varia, A Find of Isoé eae Louisiana, 186 Report of the 1970 Fern Fora 44 Rev! ye ‘the Cyathea- 88% 142; T The oe nd Fern Allies of the Sierra Nevada n California a sage evada, 44; Flora de Venezue la, Vol. I. Helechos, . Flore e Na ftalst 2 et © S Gen Polypodiacene! 95; A Manua A Monograph et Fern rs say Exiowors, 188; Pollen rs yet of Chile. is 5; Rouffa, ‘s. An Appendageless Psilotu Introduction to Aerial Shoot Me caindiog s: umohra adiantiformis, 96 aren imbricata, 9 191 Schemochromic Blue Leaf-surfaces of Selagi- nella, ale Schimeacen Schizoloma, a abe oe Sela canes 137, 138, 149, » 154; ser Arenicolae, 149; arizonica, 150, 151, 153; asprella, 149 ; bigelovii, 149, 153, 154; ser. Circinatae, , 151; cinerascens, 149- 151, 154, 157; cuspidata, 160; eremophila, 150, 151, 157, 158; ser. Eremophilae, 151; lepidophylla, eee + 157,. 159, 160; allescens, Rice 151% ; ser. Rupes- tres, 149; r. Sartori 151; subg. Selagi- nella 149; pty sect. Toon 149; 151; willd enovii. Daag aprercel Sen, T. & U. fen” iN sata Variety of Lindsaea odorata Roxb. from Darjeeling in the Himalayas, en, U. (see T. Sen) Sharp, A. J. H Webster, Jamesonia as alstonii in Oaxaca, Mexico, Smith, A. R. The Thelypteris normalis — in the Southeastern United Sta Sphaeropteris, 142 , 55, 57; chusana, 57 Stenochlaena, 119, 122; areolaris, 120, 122; 20, 122: fraxinifolia, 121; juglandifolia, 119, 121; laurifolia, 121, 122; mildbraedii, 120, 121; milnei, 120-122; palustris, 1 enuifolia, 120 Stromatopteris, 83; oe ial A Study of Azolla pinnata R. Bro Sweet, A. & L ls A pee Re pee pinnata R. B Syngramma, 41 Tectaria, 119, 145, 147, 148; blumeana, 148; irregularis, 148; dolichosora, 147, 148; te) 7 Teratoph Thelypteridaceae, 183 Thelypteris, 21, jen nt 1 80; augescens, 28— 31; cens is, 31; augescens X ma ovata, 31; ¢ vicaarsarpg 20, var. enc dioides, 20, ‘ goyazensis, 20; ntata, 22, 26, hapa eres ; Suyan 17-20; ‘fe. ubg en- iscium, 17; alis, 21, 23, 31; normalis X< ovata, 31; ovata, 27-30, var. ovata, 23, , 29, 31, var. lindheimeri, 23, 30, 31; palustris, 172, 174; patens, 26, 28, var. patens, 22, 26; puberula, 29; quadrangu laris, 24, 8 var. quadrangularis, 25, 26, 28, var versicolor, 22, 23, 31; reducta, rra, 29; torresiana, 43; totta, a1, ais hirsuta, 22. 192 var. totta, 22; uliginosa, 43; unca, 27, 28; X versicolor, 25, 26 The Thelypteris normalis Complex in the Southeastern Uni sie ‘sacs Thomas, ous Adder’s-tongue a in Louis ana, 39 Tm is, 82; tan is, 83 Trichiters 142, 168: nesotin, 166, 168 ta ic re eer chianu: acisaige ig steel oe _ Tr ryon , R. M., Jr. The ne of the ‘Two. Nev F Rica, 166 Va reschi, - Flo ie ae = Tuk i Helechos (rev ~eo The pearl ee in Spore = and Germination in Dryopteris Tax: Vascularization of =n Fan 87 AMERICAN FERN JOURNAL Vittariaceae, 41, Votava, F. ee riage T. Mic Wagner t Lindsee ET a eanifolik ‘Swart in Hawa ee D. . ee harp) W. H. Wagner, Jr. The pore Size and Germination : e Genus Selaginella in Baja California, Mexico, 14 ae aes 90; peers 32, 47, 88, 89; scopulina, Pcl 174-180; oe 172, 174 A is ngly Localized Spec of Pyrrosia, Ply C. H., Jr. (see R. C. Lommasson) Errata Page 30, line 14: For ‘‘Lindheimer” Page 46, line 1: For ‘““MoNnTGONERY” read ‘‘Lindheimert.’”’ read MONTGOMERY. Page 53, line 9: For “‘enszfolta”’ read ‘‘ensifolium.’ Exotic and Hardy Ferns 2131 Phone $63-2898—Area Code 707 Open Saturdays and Sundays from 10 A.M. to BOLDUC’S GREENHILL NURSERY Vallejo Street St. Helena, California 94574 4 P.M. and by appointment Mail orders accepted Statement of Ownership, Management and Circulation In accordance with the rules and regulations of the United States Post Office, as established under a act of October 23, 1962, Section 4369, Title ublished. 39, United States Code, the following statements are pu Title: American Fern JOURNAL Frequency of Issue: Quarterly aan a March 31, June 30, September 30, and Decem Location of Office of Publication eink: 3110 Elm Avenue, Baltimore, Maryland 21211 Location of Business Office of Publishers (Not Printers): Dr. LeRoy K. Henry, Section of Plants, Carnegie Museum, Pittsburg, Pa, 15213 Publisher: American Fern Socrety, Inc., Section of Plants, Carnegie Museum, Pittsburg, Pa. 15213 Editor: Dr. David B. Lellinger, Department of Botany, Smith- ian Institution, Washington, D. C. 20560 Managing Editor: None. Owner: American Fern Society, Inc., Section of Plants, Carnegie Museum, Pittsburgh, Pa. 15213 Mortgagees and other Security Holders: None Average No. each — an issue d pre- ceding 12 months ing date 1400 Total number of copies printed: Paid circulation 1. Sales through dealers, etc. 0 0 2. Mail subscriptions 951 953 Total paid circulation 951 953 Free distribution, including samples 0 0 Total distribution 951 953 Office use, left-over, etc. 449 447 Total 1400 1400 The statements made above are certified to be correct, and are signed by: David B. Lellinger, editor-in-chief of the American Fern Journal on Septem-— ber 28, 1971. TRIARCH INCORPORATED -A Service to Biologists = To better serve the biological community: 1. TRIARCH produces and stocks nearly 3000 dif- ferent prepared microscope slide items for use in gen- eral botany, plant anatomy, plant physiology, phyto- pathology, bacteriology, general zoology, vertebrate histology, vertebrate embryology, and parasitology. 2. TRIARCH offers to prepare slides of new or spe- cial items for a nominal fee or in exchange for pre- served materials. 3. TRIARCH advertises in biological journals to help support the financial needs of the associated So- ciety. For our current catalog No. 15, or to request special service, write to: Paul L. Conant, President TRIARCH INCORPORATED Ripon, Wisconsin 54971